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Do Not Take From This Room 





of the 

A manual for the identification of the more 

common marine invertebrates, compiled 

by Ralph I. Smith with the assistance 

of many other contributors 





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Contribution No. 11 

Systematics-Ecology Program, Marine Biological Laboratory 

Woods Hole, Massachusetts 



Copyright 1964 by the Marine Biological Laboratory 
Tenth Edition 

Referrals To 
Marine Biological Laboratory 

Supply Dept. 
Woods Hole, Massachusetts 02543 

Printed by Spaulding Co., Inc., 29 Pacella Park Drive, Randolph, Mass. 02368 



List of Plates v 

Foreword vi 

Editor's Preface vii 


I . Phylum PORIFERA 1 

Glossary 1 

Key 2 

Checklist 6 

II. Phylum CNIDARIA, Class Hydrozoa 8 

Glossary 8 

Key 10 

Checklist 17 

III. Phylum CNIDARIA, Class Scyphozoa 21 

Key to attached Scyphozoa 21 

Key to Scyphomedusae 21 

IV. Phylum CNIDARIA, Class Anthozoa 25 

Key 26 

Checklist 26 

V. Phylum CTENOPHORA, key and checklist 29 


Key to common Turbellarians 30 

Checklist of Acoela, Rhabdocoela, and Alloeocoela 32 

Checklist of Tricladida 36 

Key to Polycladida 36 

Checklist of Polycladida 37 


Key 40 

Checklist 43 

VIII. Other Unsegmented Worms: Phylum ASCHELMINTHES 46 


IX. Phylum ANNELIDA 47 

Classes Hirudinea, Oligochaeta, and Archiannelida 47 

Polychaete Terminology 48 

Key to Polychaete Families 51 

Keys and checklists of Polychaete Species 63 

X. Phylum ARTHROPODA, Subphylum Chelicerata 84 

Xiphosurida 84 

Key to Pycnogonida 84 

Checklist of Pycnogonida 85 

XI. Phylum ARTHROPODA, Lower Crustacea and Cirripedia 88 

Cephalocarida, Branchiopoda, Ostracoda, Mystacocarida, 

Copepoda , Branchiura 88 

Key to Cirripedia 89 

Checklist of Cirripedia 91 

Rhizocephala 92 

iv Table of Contents 

XII. Phylum ARTHROPODA, Peracaridan Crustaceans 93 

Key to Orders of Peracaridans 93 

Part 1. Mysidacea 93 

Key 94 

Checklist 94 

Part 2. Cumacea 98 

Key 98 

Checklist 99 

Part 3. Isopoda and Tanaidacea 102 

Key 102 

Checklist 105 

Part 4. Amphipoda 107 

Keys to Suborders 112 

Key to Families of Gammaridea 112 

Checklist of Gammaridea 114 

XIII. Phylum ARTHROPODA, Decapod and Stomatopod Crustaceans 117 

Key to Decapods 117 

Checklist of Decapods 121 

Order Hoplocarida (Stomatopoda) 127 

XIV. Phylum MOLLUSCA, Introduction 129 

Outline classification 130 

Glossary 131 

Classes Amphineura and Cephalopoda 133 

Key to Shelled Gastropoda 134 

Checklist of Shelled Gastropoda 139 

Key to Bivalvia 146 

Checklist of Bivalvia 150 

XV. Phylum MOLLUSCA, Shell-less Opisthobranchia 153 

Key 154 

Checklist 162 



Glossary 167 

Key 170 


Asteroidea, key 188 

Ophiuroidea, key 188 

Echinoidea, key 188 

Holothuroidea, key 189 

Asteroidea, checklist 189 

Ophiuroidea, checklist 189 

Echinoidea, checklist 192 

Holothuroidea, checklist 192 

XIX. Protochordates 193 


Phylum CHORDATA, Subphylum Urochordata 193 

Key to Ascideans 194 

Checklist of Ascideans 198 

Glossary of Common Names 200 

Systematic Index (Genera and Higher Taxa) 202 



Plate Pa g e 

1. Porifera 5 

2. Hydrozoa 13 

3. Scyphozoa and Anthozoa 23 

4. Turbellaria (except Polycladida) 35 

5. Polycladida, Nemertea, Sipunculoidea, Nectonema 45 

6. Polychaete Setae 51 

7. Polychaeta (1) and Archiannelida 53 

8. Polychaeta (2) 55 

9. Polychaeta (3) 57 

10. Polychaeta (4) 59 

11. Pycnogonida; Cirripedia 87 

12. Mysidacea 97 

13. Cumacea 101 

14. Isopoda (1) and Tanaidacea 109 

15. Isopoda (2) ; Hyperiidea and Caprellidea Ill 

16. Decapod Crustacea (1) 123 

17. Decapod Crustacea (2) and Hoplocarida 125 

18. Mollusca (1) 143 

19. Mollusca (2) 145 

20. Shell-less Opisthobranchia (1) 157 

21. Shell-less Opisthobranchia (2) 159 

22. Shell-less Opisthobranchia (3) 161 

23. Ectoprocta (1) and Entoprocta 181 

24. Ectoprocta (2) 183 

25. Ectoprocta (3) 185 

26. Ectoprocta (4) 187 

27. Echinodermata 191 

28. Protochordates 197 


Cape Cod juts eastward over the continental shelf as the flexed arm of Massa- 
chusetts, partly sheltered on its southern side by Monomoy, Nantucket, Martha's 
Vineyard, and the Elizabeth Islands. The Cape, islands, eastern shores of the state 
bordering the Cape, Cape Cod Bay, Nantucket Sound, Vineyard Sound, and Buzzards Bay 
constitute the Cape Cod Region. Glacial in origin, the Region is dotted by myriad 
small embayments and estuaries, and the clean tidal waters of the North Atlantic 

wash its shores a diverse ecologic setting accounting for the richness of the 

local marine biota, and for establishment of the Marine Biological Laboratory here 
in 1888. 

On September 1, 1962, the Marine Biological Laboratory inaugurated the Syste- 
matics -Ecology Program and embarked on a basic, year-round, long-term program of 
research and training in marine systematics and ecology. The Program has as one 
of its major objectives the gradual extension, in a broad sense and at a high lev- 
el of excellence, of systematic and ecologic knowledge of organisms in the estuar- 
ine and marine waters and bottoms of the Cape Cod Region. 

Early in the life of the Program we realized the lack of and recognized the 
urgent necessity for illustrated, referenced, indexed sets of keys and check lists 
of the commoner plants and animals of the Region for use by non-systematists in the 
Woods Hole scientific community and neighboring areas. A start on this task was 
provided in 1960 when Dr. Ralph I. Smith and others of the MBL Invertebrate Zoology 
Course staff assembled a small loose-leaf collection of keys then existing in the 
laboratory instructions of that course. These early keys had evolved over a period 
of years, and the authorship of some could not be determined. 

It was a happy occasion for us when Ralph Smith agreed to compile and edit 
this present publication. He arrived in Woods Hole early in the summer of 1963, 
worked indefatigably with us during that summer, then returned to Berkeley where 
he continued his labors, somehow sandwiched among his many duties at the Univer- 
sity. Now, some ten months later, the manuscript is ready for publication. We 
extend to Ralph our hearty thanks! A most sincere word of appreciation is also 
due collaborators without whose excellent cooperation and zeal the manuscript 
would never have been completed, and Mrs. San Lineaweaver for her patient and care- 
ful typing and correcting of the master copy. The preparation of this publica- 
tion •••as made possible by support of the Ford Foundation to the Systematics Di- 
vision of the Systematics-Ecology Program; it is a pleasure to acknowledge this 

The first edition of the Invertebrate Keys is a beginning. We welcome sug- 
gestions, and look forward to producing revised editions as the invertebrates of 
the Region become better known. We are also planning to produce similar keys to 
other organisms of the Woods Hole Region as this becomes possible. 

Melbourne R. Carriker, Director 
Victor A. Zullo, Resident Systematist 

Systematics-Ecology Program 
Marine Biological Laboratory 
May, 1964 Woods Hole, Massachusetts 


This handbook of keys is an attempt to fill an obvious need at Woods Hole, the 
need for a general reference on marine invertebrates for the use of students and in- 
vestigators who want to know what is here and how to identify it. 

Relatively few present day biologists realize the difficulties involved in iden- 
tifying with certainty the myriad species of marine invertebrates. Some species are 
readily recognized by distinctive form or color, but in most cases the beginner will 
experience difficulty in making a positive identification, and in many instances the 
assistance of a specialist may be necessary. Keys are useful mainly in the identi- 
fication of common and obvious animals, or in making preliminary identification of 
less common animals (perhaps to family or genus) , or by indicating to the observant 
non-systematist that he has something out of the ordinary which should be referred 
to a specialist. The problem is akin to that of medical diagnosis; the alert zo- 
ologist, like the general practitioner, need feel no shame if he makes some identi- 
fications with confidence, but seeks consultation on others. 

The purpose of zoological classification is to arrange animals into groups on 
the basis of fundamental similarities. The completion of this task will presumably 
reveal evolutionary relationships, but the immediate objective of a set of keys is 
more utilitarian, that of identifying and providing the names of the local inverte- 
brate fauna as simply, accurately, and painlessly as possible. It cannot be too 
strongly emphasized that keys are shortcuts and often very misleading; their func- 
tion is merely to clear the way to an approximation, and identifications made by 
keys, if to be of scientific reliability, should be checked by reference to the 
original description or to recent monographs. A key is made to cover a selected 
list of species, and any species not one of that group will not "key out" or, worse 
yet, may key out as something which it is not, hence the need for a cross check in 
all cases of doubt. 

The terminology of systematists may be difficult for the non-systematist to 
grasp, owing to the independent evolution of descriptive jargon within the groups 
of specialists on different animal phyla. Therefore each key will when necessary 
be accompanied by a glossary and illustrations of the descriptive terms and by a 
brief statement of the characteristics used in the taxonomy of that particular group 
and how the student may locate and recognize them. The spicules of sponges, the se- 
tae of polychaetes, the ossicles of sea cucumbers, are of basic importance to sys- 
tematists in establishing the degrees of similarity which underlie classification; 
the problem of the biological role of these structures as functional parts of the 
animals concerned lies outside the immediate field of classification. 

An aspect of identification of animals that commonly proves annoying and frus- 
trating to experimental biologists is the "name-changing" that seems so often to 
occur in systematics, resulting in the production of synonyms for some of the com- 
monly used experimental animals. In the annotated check lists we have attempted 
to explain the more conspicuous synonymies. Experimentalists should realize that, 
in an active field, terminology must change (it is no longer sufficient to speak 
merely of "Vitamin B", and such terms as muriatic acid, yellow enzyme and even DPN, 
while understandable in context are considered outdated today). In the same sense 
it is not good current usage to apply such well-known older names as Nereis lim - 
bata and Nassa obsoleta , although these particular names are perfectly understand- 
able when encountered in the older literature. It is hoped that this manual will 
provide useful information on the synonymies of local experimentally used animals 
and contribute to a better understanding of the efforts of systematists to provide 
a more rational and stable nomenclature. The short discussion of nomenclatorial 
rules in Hyman ' s "The Invertebrates", Vol. I, 1940, pages 22-26 will be found use- 
ful to those who wish to review this topic briefly, and the more extended discus- 
sion in Mayr , Linsley and Usinger's "Methods and Principles of Systematic Zoology", 
Mr-Graw-Hill (1953) provides a more complete account. In case there are any labora- 
tory biologists who feel that animal classification is outside their proper sphere 

viii Editor's Preface 

of interests, we recommend "A Classification of Living Animals", Wiley (1961), by 
no less an experimentalist than Lord Rothschild. And for those who decide to plunge 
into serious systematic work, two more references must be mentioned: "International 
Code of Zoological Nomenclature", published by the International Trust for Zoologi- 
cal Nomenclature, London (1961); and Schenk and McMasters, "Procedure in Taxonomy", 
third edition, Stanford University Press (1956). 

This manual does not pretend to be an exhaustive faunal survey of the Woods Hole 
Region which, for our purposes, is that area in which one may reasonably collect, us- 
ing Woods Hole as a base. Hence it includes not only the animals of Buzzards Bay 
and Vineyard Sound, but also species commonly taken on the northern shore of Cape 
Cod, as at Barnstable. Only by limiting its scope and content can the manual be 
kept simple and inexpensive enough to be useful and available to the students and 
beginning investigators who need it most. There are in existence two old but com- 
prehensive faunal surveys which contain much valuable information for the field 
worker : 

Verrill, A. E. and S. I. Smith, 1873. Report upon the invertebrate animals 
of Vineyard Sound and the adjacent waters. Rept. U. S. Fish Comm. 1871- 
1872: 295-778 (also published as a separate volume in 1874). 

Sumner, F. B., R. C. Osburn, and L. J. Cole, 1913. A biological survey of 
the waters of Woods Hole and vicinity. Section III. A catalogue of the 
marine fauna. Bull. Bur. Fisheries, 3_1: 545-794. 

A work much used at Woods Hole in recent years is Miner's "Field Book of Seashore 
Life", Putnam (1950). This is illustrated and has much textual material, but since 
it covers the wide area from Laborador to Cape Hatteras and is not provided with 
diagnostic keys, it is inadequate for critical identification of many local species 
and its terminology is not up to date. 

The present keys will be found to vary in ease of use and in completeness of 
coverage, and will naturally become less reliable with increasing distance from 
Woods Hole. Some groups, e.g. amphipods , are still too incompletely known to per- 
mit the construction of specific keys that would be generally useful. 

For certain important groups which are intrinsically difficult because of the 
number of species and small size of individuals, such as Protozoa, nematodes, os- 
tracods, no keys have been attempted. Those wishing to embark upon studies of such 
groups will need special instruction and more detailed literature than we can pro- 
vide. Plankton, especially its component of immature stages, will not be treated 
in these keys, although representatives of certain groups may be identifiable. In 
other instances, the keys will serve not to provide answers, but to clarify the 
problem. We regard this edition of the keys as tentative: they will be found inac- 
curate or incomplete in many respects. We should appreciate receiving criticisms 
or suggested revisions at any level. 

In Woods Hole at the present time the preponderance of experimentalists places 
a burden of identification upon those who are undertaking systematic problems and 
who are expected to make reliable identifications. With the aid of proper keys, 
this load can be more fairly distributed, routine identifications accomplished, non- 
identifiable animals more readily detected and referred to specialists, and the ex- 
change of vital information between laboratory and field workers facilitated. 

This manual has been made possible only by the generous assistance and coopera- 
tion of numerous individuals, actually too numerous to list fully. However, cer- 
tain ones have given very liberally of help and advice, and are listed here, to- 
gether with an indication of the areas in which they contributed. It should be 
noted that there is no clear distinction between those listed as "authors" of chap- 
ters and those who contributed heavily to chapters in which a good deal of editorial 

Editor's Preface ix 

compilation has taken place. It seems appropriate to acknowledge gratefully this 
help, and to express the wish that users of this manual will feel free to pass on to 
editors, and advisors alike, the criticisms and comments that will be inevitable. 

Porifera: Willard D. Hartman , Peabody Museum of Natural History, 
Yale University, New Haven, Connecticut. 

Coelenterata (Hydrozoa) : Sears Crowe 11 , Department of Zoology, 
University of Indiana, Bloomington, Indiana. 

Coelenterata (Hydrozoa, Stauromedusae) : G. F. Gwilliam , Department of Biology, 
Reed College, Portland, Oregon. 

Coelenterata (Hydrozoa, Stauromedusae): Kay Werner Petersen , Systematics-Ecology 
Program, Marine Biological Laboratory, Woods Hole, Massachusetts. 

Coelenterata (Anthozoa) : Cadet Hand , Department of Zoology, 
University of California, Berkeley, California. 

Platyhelminthes : Louise Bush, Department of Zoology, 
Drew University, Madison, New Jersey. 

Annelida (Polychaeta) : Marian H. Pettibone , Division of Marine Invertebrates, 
U. S. National Museum, Washington, D. C. 

Pycnogonida: Joel W. Hedgpeth , Pacific Marine Station, 

University of the Pacific, Dillon Beach, Marin County, California. 

Crustacea (Cirripedia) : Victor A. Zullo , Systematics-Ecology Program 
Marine Biological Laboratory, Woods Hole, Massachusetts. 

Crustacea (Mysidacea, Cumacea, Isopoda) : Roland L. Wig ley , Biological Laboratory, 
U. S. Bureau of Commercial Fisheries, Woods Hole, Massachusetts. 

Crustacea (Isopoda): Thomas E. Bowman , Division of Marine Invertebrates, 
U. S. National Museum, Washington, D. c 

Crustacea (Amphipoda) : Eric L. Mills , Department of Biology, 
Queen's University, Kingston, Ontario, Canada. 

E. L. Bousf ield , Natural History Branch, 
National Museum of Canada, Ottawa, Canada. 

Crustacea (Cephalocarida) : Howard L. Sanders , Biology Department, 

Woods Hole Oceanographic Institution, Woods Hole, Massachusetts. 

Mollusca (General) : W. Russell Hunter , Department of Zoology, 
Syracuse University, Syracuse, New York. 

Stephen C. Brown 

709 Arch Street, Ann Arbor, Michigan. 

Mollusca (Shell-less Ophisthobranchia) : George M. Moore , Department of Zoology, 
University of New Hampshire, Durham, New Hampshire. 

Entoprocta and Ectoprocta: Mary D. Rogick , Department of Biology, 
College of New Rochelle, New Rochelle, New York. 

Editor's Preface 

Echinodermata: John M. Anderson , Department of Zoology, 
Cornell University, Ithaca, New York. 

Urochordata and Mystacocarida: Donald J. Zinn , Department of Zoology, 
University of Rhode Island, Kingston, Rhode Island. 

Urochordata: Donald P. Abbott, Hopkins Marine Station, 

Stanford University, Pacific Grove, California. 

Compiling and editing: Ra lph I . Smith , Department of Zoology, 
University of California, Berkeley, California. 

Production of the manual: Melbourne R. Carriker , Systematics-Ecology Program, 
Marine Biological Laboratory, Woods Hole, Massachusetts. 

Artists: Mrs . Ruth L. von Arx , Mrs . Emily Read , Bruce Shearer , Ann H. Miller 
(cover design), and several of the above contributors. 

My thanks are due the staff and students of the MBL Invertebrate Zoology Class 
of 1963, who cooperated extensively in collecting material and critically testing 
keys. The Supply Department has also been most helpful, and I owe much to Mr. John 
Valois for his help in the summer of 1963, and especially I am indebted to Mr. Mil- 
ton Gray, as whose helper in 1939 I learned much that has proved useful in all my 
work in marine zoology. Above all, the unstinting support given to the prepara- 
tion of these keys by the Systematics-Ecology Program of the MBL has been the de- 
cisive factor in their publication at this time. I wish to record my appreciation 
for all that Dr. Melbourne R. Carriker and his entire group have done to facili- 
tate this task, and especially am I grateful to Mrs. San Lineaweaver and Mrs. Vir- 
ginia Smith for their extraordinary efficiency and cheerfulness in the typing in- 

Ralph I. Smith, Editor 
May, 1964 

Chapter I ] 


By Willard D. Hartman 

There are about ten common sponges in the Woods Hole region. Some are identifi- 
able on sight by form and color; others must be carefully examined before an identi- 
fication is ventured. Miner's "Field Book" contains inaccuracies and is almost use- 
less for identification of sponges. De Laubenfels (1949) has a selection on methods 
of handling and observing sponges. Hartman (1958) has given a systematic account of 
southern New England sponges together with notes on their natural history. 

Some knowledge of the terminology of sponge spicules is neccessary if the taxo- 
nomic literature on the group is to be read. In the following glossary, terms com- 
monly used are defined to aid those who may go further in this field, as well as to 
make clear certain expressions used in the key. Most of these terms are illustrated 
in Plate 1. 


Acantho -: prefix meaning spined. 

Aniso -: prefix meaning unequal. 

Anisochela : a chela with unequal ends. 

Chela : a microsclere type resembling a pair of anchor flukes or a shovel with a blade 

at each end, the ends bent toward each other. 
Isochela : a chela with equal ends. 
Megascleres : the larger spicule types, comprising the major structural elements of a 

sponge skeleton. 
Microscleres : relatively very small spicules of various form, cVij.-acteristic of cer- 
tain sponges. 
Monaxon ; an unbranched spicule. 

Oxea : a monaxon spicule (megasclere) pointed at both ends, often slightly curved. 
Sigma : a microsclere of C- or S- shape. 
Spiraster : angulate rod-shaped microscleres provided with spines which may be aligned 

in a spiral course arour.d the rod. 
Sponqin : proteinaceous horny skeletal material, either alone (as in bath sponges), or 

binding siliceous spicu es together (as in Haliclona or Microciona ) . 
Style : a monaxon spicule wi :h one end pointed, other end blunt. 
Subtylostyle ; a monaxon spicule with one end pointed and the other with an indistinct 

swelling or knob (halfw y between tylostyle and style) . 
Toxon : a bow-shaped microscvere (pi. toxa) . 
Triaene a spicule with 4 rays, one of which is much longer than the other 3 (known as 

clads) . Anatriaenes have the clads directed backward; protriaenes have the clads 

directed forward. 
Tylo : a prefix meaning a rounded or ball-shaped structure. 

Tylote : a monaxon spicule with rounded knobs on both ends (e.g., in Lissodendoryx ) 
Tylostyle : a style with the blunt end in form of a little swelling or knob (e.g. in 

Cliona ) . 

A Note on Microtechnique 

Detailed studies of sponge anatomy may require the preparation of microscopic sec- 
tions. For gross preservation 95% alcohol or neutralized formalin followed by 75% al- 
cohol can be used. Neutral formalin is best prepared by adding hexamethylenetetramine 
(= hexamine) to commercial formalin (37 to 40% formaldehyde) in the proportion of one 
pound of hexamine to one gallon of formalin. This stock solution is then diluted with 
water (9 parts water; 1 part neutral formalin) to give 10% formalin or 4% formaldehyde. 
Formalin that has not been neutralized, or that has been neutralized with borax, should 
be avoided for preserving sponges since both may cause maceration. When formalin neutra- 
lized with hexamine is used, the sponges should be transferred to 75% alcohol after a 
few days. Fixation of small pieces of each species in standard histological fixatives 

2 Porifera 

is desirable, as often histological work must follow. Thin, free-hand sections are 
readily made with a razor blade if the specimen has been hardened in alcohol. Sections 
perpendicular to the surface of the colony as well as tangential sections are useful. 
Such sections can be directly dehydrated, cleared in xylol, and mounted in piccolyte 
or damar or they can be stained with a saturated solution of basic fuchsin of safra- 
nin in 95% alcohol before clearing and mounting. In studying sections, the various 
types of spicules can be observed as well as their arrangement in the skeleton. Other 
structures to look for are a cortex, often packed with spicules of one category, sub- 
dermal spaces, tracts of spicules, and spongin fibers with or without enclosed spi- 
cules. In some cases the shapes and dimensions of the flagellated chambers are im- 

In sections, certain spicule categories may be overlooked, and it is usually de- 
sirable to make slides which have been freed from the cellular elements of the colony. 
This can be done by treating a small part of the specimen with sodium hypochlorite 
(clorox will serve) until the spicules are free. For temporary mounts a piece of 
the sponge can be placed on a microscope slide with a drop or two of clorox. After 
disintegration of the cells a cover slip is added and the spicules are ready for ex- 
amination. If permanent mounts are desired, a piece of sponge can be placed in a 
centrifuge tube along with a snail amount of clorox. After disintegration of the 
cells, the sample is then washed several times with water, centrifuging between wash- 
ings to make certain that the minute microscleres have settled. Finally the water 
can be replaced by 95% alcohol, and the suspension of the spicules poured onto a 
slide and allowed to dry on a warming plate (or the alcohol can be ignited). A drop 
of xylene is added to the dry preparation, followed by piccolyte or damar and a co- 
ver slip. A slide of clean spicules is now available for study and measurement of 
spicule types. 


(Figure references are to Plate 1) 

1. Structure of sponge simple; tubular to urn-shaped; pale tan 

to whitish; spicules calcareous (test with 10% acetic acid under coverslip) .... 2 

1. Structure massive, fleshy, or spongy; encrusting or branching; 

color various; spicules siliceous 3 

2. Sponges in form of branching, cylindrical, pale tubes (fig. 1) 

Leucosolenia sp. 

2. Sponges in form of little urns or vases, usually clustered, 
with fringe of spicules around the terminal osculum (fig. 2) 

Scvpha sp. 

3. Sponges obviously boring into and sometimes overgrowing shells 
and other calcareous material; color brilliant sulfur -yellow 

to pale yellow Cliona 4 

3. Sponges not boring; form and color various 7 

4. Sponge brilliant sulfur-yellow, boring into calcareous sub- 
strate or massive and free-living; external perforations in 
substrate large (incurrent, 0.8-2.5 mm across; excurrent, 

2.0-4.5 mm across); tylostyles only (fig. 31 Cliona celata 

4. Sponge light or pale yellow, always boring into calcareous 
substrate; external perforations small (incurrent, 0.2-0.5 
mm across; excurrent, 0.6-1.6 mm across); microscleres pre- 
sent in addition to tylostyles 5 

5. Spirasters only as microscleres (fig. 4) Cliona lobata 

5. Spined or smooth oxeas present as well as spirasters 6 

Porif era 

6. Spirasters usually angulate and larger (6-23n x 1.0-3.8u) 

(fig. 5) ; external perforations in substrate often arranged 

in circular patterns Cliona vastifica 

6. Spirasters less distinctly angulate and smaller (6-15(i x 

0.5-3. 2a.) ; tends to occur in brackish water Cliona truitti 

7. Megascleres oxeas only 8 

7. Megascleres other than oxeas 12 

8. Megascleres smaller «200u. in length) and conspicuously 
joined together by more or less spongin to form a network 
or a system of tracts (fig. 6); dermal skeleton absent 

Haliclona 9 

8. Megascleres larger (mean length > 200|i) , arranged in loose 

tracts with little spongin or occurring at random; distinct 

dermal skeleton present (figs. 9, 10) Halichondria 11 

Form branching into rounded or flattened fingers arising 
from a narrow stalk; oscules distributed along sides of 
branches; color yellowish to tan, often tinged with violet 

Form encrusting, often with low tubules arising from a 
basal encrusting mass 

Haliclona oculata 




Sponge a flat encrustation; oscules not raised on tubules; 
excurrent channels converge on oscules to provide conspic- 
uous stellate patterns beneath surface; gemmules absent; 

color light buff or tan to brown or gold Haliclona canaliculata 

Sponge an encrustation from which arise numerous low tubules 
often bearing terminal oscules, consistency very soft; white 
gemmules form at base of sponge in late summer; color dark 
tan to gold, often tinged with pink or lavender (figs. 6-8) 
Haliclona loosanof f i 






Sponge encrusting and provided with numerous, low, upright 
tubules each terminating in an oscule; dermal skeleton 
tends to be a regular network of multispicular tracts (fig. 

9) ; color orange-yellow to greenish Halichondria panicea 

Sponge encrusting to massive, usually provided with upright 
tubules which may bear terminal oscules; some colonies con- 
sist of masses of anastomosing branches; dermal skeleton 
tends to consist of a less regular network of multispicular 
tracts with larger meshes (fig. 10); color orange-brown to 
yellow-beige to olive-green 
Halichondria bowerbanki 

Color bright red to orange-brown; sponge encrusting to com- 
plexly branched; spicules include styles to subtylostyles 
with spined or smooth heads, short acanthostyles, toxa, and 

isochelas (fig. 11) Microciona prolifera 

Color never bright red; spiculation otherwise 13 

Megascleres are tylostyles 14 

Megascleres are otherwise 16 



Microscleres absent; sponge a thin encrustation on rocks 
and algae; basal layer of tylostyles arranged with heads 
against substrate; color orange-brown, hazel, or olive- 
brown Prosuberites epiphytum 

Microscleres present 15 

Plate 1 


Figs. 1,2,3,8 by Bruce Shearer; rest redrawn by 
Mrs. Emily Reid after Hartman (H) ; fig. 13 after 
Wilson. Scales as indicated on plate. 

Fig. 1. Leucosolenia sp., fragment of a colony. 

2. Scypha sp., three individuals. 

3. Cliona celata , colony overgrowing a Mercenaria shell. 

4. Cliona lobata : A. Tylostyles, B. Spirasters with spines (H) . 

5. Cliona vastifica : A. Tylostyles, B. Smooth and microspined 
oxeas, C. Spirasters and a microspined style. 

6. Haliclona loosanoffi , spicule tracts joined by spongin (H) . 

7. Haliclona loosanoffi , spicules: oxeas, styles, strongyles (h) 

8. Haliclona loosanoffi , specimen growing on shell of living 
Mytilus . 

9. Halichondria panicea , dermis, with spicules and pores (H) . 

10. Halichondria bowerbanki , dermis, with spicules and pores 

11. Microciona prolifera : A. Megascleres, B. Toxa, C. Isochelas 

12. Suberites ficus ; A. Styles and tylostyles, B. Microscler- 
es (H) . 

13. Mycale fibrexilis ; A. Megascleres tylostyles, B. Micro- 
scleres: sigmas, and anisochelas (after Wilson). 

14. Lissodendoryx isodictyalis : A. Megascleres: styles and ty- 
lates, B. Microscleres: sigmas, C. Microscleres : isoche- 
las (H) . 

15. Isodictya deichmannae : A. Megascleres: styles, strongyle, 
centrotylote oxea, B. Microscleres: Isochelas (H) . 


ca Imm 

ca 10 mm 

flft ft 




6 Porifera 

15. Sponge massive, lamellate or lobate; microscleres small, 

rod shaped, with central swelling, spined or smooth, abun- 
dant near surface (fig. 12); color yellow to yellowish- 
gray Suberites f icus 

15. Sponge encrusting to massive; microscleres include sig- 
mas, toxa, and anisochelas (fig. 13); color yellow-ochre, 
hazel-brown, olive-brown ranging to gray-tan, olive-gray 

to slate -gray Mycale fibrexilis 

16. Megascleres are long oxeas and triaenes ; microscleres are 
microspined sigmaspires; sponge biscuit shaped with termi- 
nal osculum and flattened base made up of matted oxeas and 

anatriaenes Craniella gravida 

16. Styloid megascleres present 17 

17. Sponge encrusting to massive; color buff to olive-tan to 
gray-brown, slate gray, or bluish-gray; dermal tylotes 
present; microscleres are sigmas and isochelas (fig. 14) 

Lissodendorvx isodictyalis 

17. Sponge with upright flattened branches; oscules distri- 
buted along sides of branches; skeleton a reticulation 
of tracts of styles; microscleres are isochelas only 
(fig. 15) Isodictya deichmannae 

Class Calcarea 

Leucosolenia sp. An "asconoid" calcareous sponge, especially valuable in the labor- 
atory. The specific identity of the common Leucosolenia at Woods Hole is at 
present uncertain. 

Scypha sp. A "syconoid" calcareous sponge. Although often called (and sold as) 

" Grantia " , it does not have the distinct dermal cortex and incurrent canals of 
Grantia . The Woods Hole species is not referable to the species lingua described 
by Haeckel (1872) from Newfoundland and needs a new name. Scypha is a prior 
name for Sycon , which is still used by many zoologists. 

Class Demospongiae 
Order Haplosclerida 

Haliclona canaliculata Hartman, 1958. Not yet recorded from the Woods Hole region 

but to be expected as an encrustation on the lower surface of rocks. 
Haliclona loosanof f i Hartman, 1958. Collected in run-out channel under bridge at 

Barnstable Harbor; records from Nonamesset Island and New Bedford area in the 

Gray Museum. 
Haliclona oculata (Linnaeus, 1759) . (= Chalina oculata of past authors) . Seldom 

found intertidally ; common in offshore waters. 

Order Poecilosclerida 

I sodictya deichmannae (de Laubenfels, 1949) . (= Neosperiopsis deichmannae de L.) 
An offshore species . 

Lissodendoryx isodictyalis (Carter, 1882) . Fairly common, but in field may be con- 
fused with Halichondria . Excellent for demonstration of microscleres. 

Microciona prolifera (Ellis and Solander, 1786) . Common and easily recognized by 
color; assumes branching form in subtidal waters. Extends into somewhat brack- 
ish lagoons. 

Porifera 7 

My c ale fibrexilis (Wilson, 1891) . (= Esperella fibrexilis Wilson, 1891; - Carmia 
f ibrexilis , de Laubenfels, 1949). Common on wharf pilings in the Woods Hole 
region; often as a thin encrustation that might be confused with Lissodendoryx ; 
sometimes massive. 

Order Halichondrida 

Halichondria bowerbanki Burton, 1930. The common Halichondria in the Woods Hole 

region. Extends into brackish lagoons. 
Halichondria panicea (Pallas, 1766). Probably occurs at Woods Hole but difficult 

to distinguish from the previous species. Common north of Cape Cod. (See 

discussion in Hartman, 1958) . 

Order Hadromerida 

Cliona celata Grant, 1826. A common boring sponge which assumes a massive, free- 
living condition as it grows older. Easily distinguished from other local clio- 
nids by the large size of the openings in the calcareous substrate excavated for 
the incurrent and excurrent papillae. 

Cliona lobata Hancock, 1849. Of common occurrence on oyster shells and other cal- 
careous materials. 

Cliona truitti Old, 1941. Not recorded as yet from Woods Hole, but may occur in 
brackish waters . 

Cliona vastifica Hancock, 1849. Of common occurrence on oyster shells and other 
calcareous materials. 

Prosuberites epiphytum (Lamarck, 1816) . Not yet recorded from Woods Hole, but is 
to be expected as a thin encrustation on the lower surfaces of rocks. 

Suberites ficus (Johnston, 1842) . A common subtidal species. 

Order Choristida 

Craniella gravida (Hyatt, 1877) . Reported common on mud bottoms in deeper waters 

of Buzzards Bay. De Laubenfels' synonymy (1949) of this species with the deep- 
water North Atlantic C. crania is in error. 


De Laubenfels, M. W. , 1949. The sponges of Woods Hole and adjacent waters. Bull. 
Mus. Comp. Zool. Harvard, 103 (1) : 1-55. 

Hartman, W. D. , 1958. Natural history of the marine sponges of southern New Eng- 
land. Peabody Mus. Nat. Hist., Yale Univ. Bull., 12: 155 pp. 

Old, M. C, 1941. The taxonomy and distribution of the boring sponges (Clionidae) 
along the Atlantic Coast of North America. Chesapeake Biol. Lab. Publ., 44: 
30 pp. 

A reference for sponges found north of Cape Cod is: Procter, 1933. Porifera. In: 

Biol. Surv. Mt. Desert Region. Part V. Marine Fauna. Philadelphia, Wistar Inst.. 

8 Chapter II 


Introduction and Class Hydrozoa 

The phylum Cnidaria is a large and complex group characterized by the produc- 
tion of nematocysts, by a tubular or cup-like body of two layers (ectoderm and endo- 
or gastroderm) separated by a mesogloeal layer, with the mouth as the sole opening 
to the digestive cavity, and by enormous diversity of form and life history. The 
best general account of the group is in Hyman ' s "The Invertebrates", Vol. 1. This 
phylum is often called the Coelenterata, but it now seems best to give up the latter 
term since it is used in some texts to include both Cnidaria and Ctenophora. The 
latter group is generally considered quite distinct, and is treated as a separate 
phylum in Chapter V of this manual. 

The present chapter deals with the Class Hydrozoa, characterized in many, but 
not all, of its members by the development of both hydroid (polypoid) and medusoid 
stages or forms in their life cycles. This class is the most numerous and diverse 
of the cnidarian classes in the Woods Hole region. 

The two other cnidarian classes are the Scyphozoa (Chapter III), most of whose 
members are large medusae, but which also includes an order (Stauromedusae) of at- 
tached forms, and the Anthozoa (Chapter IV). The latter is the largest of the cni- 
darian classes, but its members are mainly tropical, including corals and a host 
of related forms. In the Woods Hole region, sea anemones make up the bulk of the 
representation of anthozoans. 


The representation of sessile Hydrozoa in the waters of the Woods Hole Region 
is extremely good. But, unfortunately, the class Hydrozoa, and in particular the 
order Hydroida, has been plagued with a curious and troublesome double taxonomy. 
Students of the benthic fauna have developed one system of families, genera, and 
species based upon the polypoid or "hydroid" stages, while workers on the plankton- 
ic community have tended to develop a system based upon the medusae. Since in num- 
erous instances polyp and medusa of a single species have received different gen- 
eric and specific names, and since in many cases the life histories are not com- 
pletely known, the problem of synonymizing these separate stages has been diffi- 
cult and will continue so. Furthermore, when the attempt is made to bring polyp 
and medusa together under one name, the rule of priority may require that a less 
well known prior name replace a familiar later name. This does not bother sys- 
tematists greatly, but is frustrating and exasperating to the non-specialist. The 
latter may take comfort, however, in the fact that progress toward a single inte- 
grated taxonomy is being made. An excellent example of enlightened systematic 
work based upon life history studies is to be seen in F. S. Russell's "The Medu- 
sae of the British Isles" (1953) . 


Adnate: One side of structure growing attached, as the hydrothecae of Sertularia . 

Annulated: Possessing a ringed appearance. 

Athecate: The hydranth is not protected by a chitinous cup or hydrotheca ; gymno- 

blastic hydroids are athecate. 
Blastostyle: Modified zooid (gonozooid) from which the medusoids are budded; may 

be protected by a surrounding theca (gonotheca or gonangium) . 
Calyptoblastea : The order of thecate hydroids, having each hydranth protected by 

a hydrotheca. 
Capitate tentacle: Short tentacle with a very distinct terminal knob studded with 

nematocysts, as in Z anclea or Pennaria . 

Hydrozoa g 

Colony: The thecate hydroid colony is always a polymorphic system, consisting of 
at least the gastrozooids (hydranths) or food-securing individuals, and the go- 
nozooids or asexual individuals which produce by budding the sexual free or 
attached medusoids. The various zooids form a unit arising from a single plan- 
ula larva and connected by stems and branches; rootlike structures attach the 
colony to the substrate. In most athecate hydroids the colony is not polymor- 
phic, since all the hydranths alike produce medusa-buds, or else the medusa- 
buds are produced on certain areas of stolon or stem. 

Dactylozooid: A mouthless polyp armed with nematocysts, and serving for protec- 
tion or to aid in food catching, as in Podocoryne or Hydractinia . 

Distal tentacles: The set of tentacles farthest from the stalk of gymnoblasts 
that possess two or more sets of tentacles. 

Fascicled: Hydroids in which several stems of the colony are closely bound to- 
gether into a bundle or fascicle, as in some species of Eudendrium 

Filiform tentacle: Long threadlike tentacle, over which the nematocysts are usu- 
ally evenly distributed. 

Gastrozooid: A nutritive polyp; a hydranth. 

Gonangium: The entire asexual reproducing individual of the hydroid colony, in 
calyptoblasteans composed of the protecting gonotheca, the blastostyle, and 
the medusoid forms which bud from the blastostyle. 

Gonophores : Sexual individuals, which in calyptoblasteans arise asexually by 

budding from the blastostyle, being either free medusae or reduced to sporo- 
sacs; in gymnoblasteans they may arise from the stolons, hydranths, pedicels, 
or stems. However, Fraser calls the blastostyle, which bears the sexual zo- 
oids, a gonophore. 

Gonotheca: The peridermal protective structure surrounding the blastostyle and 
gonophores in calyptoblasteans. 

Gonozooid: Another term applied to any individual of the colony which buds off 
sexual individuals such as free medusae or the various degrees of reduced, 
attached sporosacs. 

Gymnoblastea : The order of athecate hydroids, lacking protective thecae about 
the hydranths. 

Hydranth: The hydranth (gastrozooid) is the feeding member of the colony, having 
a terminal mouth surrounded by tentacles; may be either sessile or stalked. 

Hydrocaulus: The hydrocaulus is the main stem of that type of colony in which 
the budding of new hydranths can take place on the stalks of the older hy- 
dranths. This gives rise to erect, branched colonies. The term hydrocaulus is 
sometimes used to refer to the type of colony developed in the above way, in 
contrast to rhizocaulus , in which the new hydranths are developed from the 
stolons only. 

Hydrocladium (-ia) : Lateral branch(es) growing from the hydrocaulus. 

Hydrorhiza: The rootlike structure, which may be more or less simple or a mass 
of tangled tubes, attaching the colony to substrate. 

Hydrotheca: The peridermal, cuplike structure that surrounds the hydranth, and 
into which the hydranth may be drawn when disturbed. 

Medusoid form: Inclusive term applied to the various types of sexual zooids (go- 
nophores) ; it includes (1) free medusae with velum, tentacles, radial canals, 
and manubrium ( Bougainvillia , Podocoryne , Obelia ) ; (2) gonophores (eumedusoid) 
that resemble developing medusae but have no tentacles or other marginal struc- 
tures, and which in some forms break away but soon die, since they lack a 
mouth; however, in most cases they remain attached ( Tubularia ) ; (3) gono- 
phores (cryptomedusoid) that are still further reduced, lacking radial canals 
and other obvious medusaLd structures ( Clava ) ; and (4) the most reduced type, 
in which the sex cells ripen directly on the sides of the blastostyle ( Ser - 
tularia ) . Types 2, 3, and 4 are often called sporosacs. The condition in 
Hydra is not unlike that in Sertularia , but there is no differentiated blas- 

Nematophore: The small, highly modified zooid (dactylozooid) of protective func- 
tion which is characteristic of the family Plumulariidae (e.g . Schizotr.icha ) 

10 Hydrozoa 

Operculum: A lid closing the hydrotheca or gonotheca when the zooid is drawn in 

(Calyptoblastea) . 
Pedicel: The stalk that bears a hydrotheca. 
Proximal tentacles: The circlet of tentacles nearer the stalk in gymnoblasts that 

have two or more whorls of tentacles. 
Rhizocaulus: See discussion under hydrocaulus . 
Sessile: Hydrotheca or other structure attached directly to stem or branch of 

colony; lacking a pedicel ( Schizotricha , Sertularia ) . 
Sporosacs: Reduced gonophores that do not develop into free medusae, but remain 

attached and produce the gametes ( Clava , Hydractinia ) . 
Stolons: Tubular or rootlike processes that extend over the substrate; the 

stolons are a part of the hydrorhizal system. 
Thecate: Hydroids that possess a protective, cuplike hydrotheca which surrounds 

the hydranth, as in the Calyptoblastea. 
Zooid: A general term applied to any of the several types of individuals or "per- 
sons" of the hydroid colony. 


Modified after key and check list used by Invertebrate Zoology Course from 1942 
through 1946. The assistance of Dr. Sears Crowell, Dr. Frank Gwilliam, Dr. Kay Pe- 
tersen, and several others in this 1964 revision is gratefully acknowledged. Fig- 
ure references in this key are to Plate 2. 

1. Hydranth naked, lacking a protective cup (hydrotheca) 

into which it can be retracted (but perisarc may extend 
to base of tentacles in Bouqainvillia ) (fig. 1) 


1. Hydranth with a protective hydrotheca into which it can 
be withdrawn (but cup much reduced in Halecium) (figs. 18, 

19) Suborder CALYPTOBLASTEA 20 

2. Hydranths with filiform tentacles only 3 

2. Hydranths with distinctly capitate tentacles only 19 

2. Hydranths with both filiform and capitate tentacles (dis- 
tal scattered capitate, basal filiform whorls) ; bushy 

colony, dark perisarc, pink hydranths (fig. 2) Pennaria tiarella 

3. Filiform tentacles scattered on hydranth (Clavidae) 4 

3. Filiform tentacles in distinct whorls 6 

4. Very small (2 cm or less) pinkish growths, sparsely 
branched or with slender hydranths arising from a sto- 
lon. Free medusae arise singly on short pedicels with 

perisarc near bases of hydranths Turritopsis nutricula * 

4. Gonophores (reduced medusoids or sporosacs) in clusters 

below tentacles or as ovoid bodies on branching stems 5 

5. In very low salinity or fresh water; colony branching, 
perisarc brownish; gonophores ovoid, borne on stem be- 
low hydranths (fig. 11) Cordylophora lacustris 

5. In marine or nearly marine salinities, pink hydranths in 
clusters arising from basal stolons; clustered sporosacs 
below tentacles Clava leptostyla 

5. (Note: a rare species, the large (2-3 cm) solitary Acaulis 
primarius also has scattered filiform tentacles and sporo- 

*A form answering this general description but with sporosacs instead of free 
medusae was taken in 1963, but is as yet unidentified. 

Hydrozoa n 

6. Hydranths with a single whorl of filiform tentacles 7 

6. Hydranths large and showy, with two whorls of fili- 
form tentacles, the proximal longer than distal 15 

6. (Note: the minute pelagic hydroid Margelopsis qib - 
besi consists of a single swimming hydranth , with 

2 subequal whorls of tentacles, and bearing medusa- 
buds) . 

7. Colonies arising from a basal mat of stolons, and 
without perisarc around stems of zooids (Hydractini- 

idae) 8 

7. Colonies erect or branching, with perisarc extend- 
ing out to bases of hydranths, or even to bases of 

tentacles 10 

8. Hydrorhizal spines rough; gonophores are sporosacs, 
borne on individuals (gonozooids) with reduced ten- 
tacles; colonies common on shells occupied by her- 
mit crabs, but also on rocks, piles, etc; color 
varies from white to salmon (fig. 17) 

Hvdractinia echinata 

8. Hydrorhizal spines smooth; gonophores are free medu- 
sae, borne on gonozooids with well developed tentacles 
(to see released medusae, let colony stand overnight 

in a small container of cool sea water) 9 

9. Medusae with 8 well developed tentacles and unripe 
when liberated (fig. 15). Colony contains "spiral 
zooids" (tentacle-less defensive individuals; look 
for these within the aperture of the snail shell, 
close to the hermit crab occupant, fig. 16); color 
white to pink; generally on shells of Nassarius 
trivittatus occupied by hermit crabs. 

Podocorvne carnea 

9. Medusae with ripe gonads but rudimentary tentacles 
at time of release. No spiral zooids in colony. 
Forms a sparse colony, generally on shells of liv- 
ing Nassarius obsoletus Stylactis hooperi 

10. Perisarc ends below hydranths; hydranths with trum- 
pet shaped hypostomes; gonophores are sporosacs (fig. 
12) Eudendrium 11 

10. Perisarc continued as a thin expansion over base of 
hydranth up to tentacles; hypostome conical; free medu- 
sae liberated; at time of liberation medusae show 4 

pairs of tentacles (fig. 1) Bouqainvillia 14 

11. Main stem fascicled (fig. 12), colonies branched, bushy 

and fairly large 12 

11. Main stems not fascicled; colonies small and sparsely 

branched 13 

12. Tentacles of gonozooids (hydranths bearing gonophores) 
showing little or no reduction; male gonophores 2- or 

3 -chambered ... „ , -, . 

19 n^~~„ ■ j t, ■ Eudendrium ramosum 

LZ. Gonozooids bearing gonophores tend (at least in some to 

be reduced ("aborted") (fig. 12); male gonophores 4- or 

5 -chambered „ , ' 

Eudendrium carneum 


Plate 2 


Sources: Fraser (F) , from living specimens (S) . Scale 
bars all 1 mm, approximately. Somewhat simplified. 

Fig. 1. Bougainvillia carolinensis , with medusae; note perisarc (un- 
stippled) up to tentacles (S) . 

2. Pennaria tiarella , hydranth with gonophores (medusae with rudimen- 
tary tentacles (S) . 

3. Clytia edwardsi , hydrotheca and gonangium (F) . 

4. Clytia johnstoni , gonangium (F) . 

5. Schizotricha tenella , detail of lateral branchlet with hydrotheca 
and 4 nematophores (S) . To scale of fig. 10. 

6. Zanclea costata , hydranth (S) . 

7. Campanularia f lexuosa , short internodes, immature and ripe gonan- 
gia (S) . 

8. Campanularia calceolifera , hydranth (tentacles schematic) and male 
gonangium, long internodes (S) . 

9. Same, female gonangium. 

10. Sertularia pumila, opposite, adnate, hydrothecae (S) . 

11. Cordylophora lacustris , with young gonangium (S) . 

12. Eudendrium carneum, fascicled stem, gonophores on gonozooid with 
"aborted" tentacles (S) . 

13. Obeli a sp. , a "bottle-necked" gonangium (F) . 

14. Calycella syringa , hydrotheca with operculum (S) . 

15. Liberated medusa of Podocoryne carnea (S) . 

16. Part of colony of Podocoryne on shell used by hermit crab. Note 
medusa-buds at left, and spiral zooids at angles of shell aperture 
(those at right slightly enlarged) (S) . 

17. Individuals of Hydractinia echinata, among roughened spines. Note 
that neither the reduced tentacles of dactylozooids and gonozooids 
nor the feeding tentacles are classed as capitate (S) . 

18. Halecium halecinium,female gonangium bearing pair of hydranths (F) . 

19. Same, with gastrozooid and male gonangium (F) . 

Plate 2 



13. Tentacles of gonozooids not completely reduced; male 
gonophores 2- or 3-chambered; colony small (1/2 inch) , 
whitish Eudendrium album 

13. Male gonophores borne on fully reduced zooids; colony 

pink , 1/2 to 1 inch high Eudendrium tenue 

14. Stout bushy growths; stems fascicled; greenish peri- 
sarc, reddish hydranths; tentacles in extension long,, and seeming to point in all directions; 2- 
8 inches high; medusae budded singly from branches 
(fig. 1). This is the Bougainvillia hydroid generally 

collected Bougainvillia carolinensis 

14. Delicate sparse growth with a single whorl of filiform 
tentacles (Russell) . This species is well known by 
its medusa, but the status of the hydroid at Woods 
Hole is not clear Bougainvillia super ci liar is 

15. Zooids solitary, not commonly found; liberate free me- 

15. Zooids generally in branching form or clustered; medu- 
sa-buds not released 


16. Zooid with perisarc very weak or lacking; large (1-4 

inches) Corymorpha pendula 

16. Zooid with obvious brown perisarc; single or slightly 

branched; about 24 short distal (oral) tentacles, 30 

longer proximal tentacles; easily mistaken for Tubularia 

until medusae seen 
Ectopleura dumortieri 



Colony branching; stems extensively annulated; hydranths 
bright pink; colonies not large (1 to 1.5 inches) 

Colonies are clusters of long stems, unbranched or sparse- 
ly branched; stem annulated only at intervals 

Tubularia larynx 


18. Grows as a dense cluster of many sparsely or unbranched 
stems; hydranths rose colored; 3-5 inches high 

18. Forms clusters of unbranched stems up to 7 inches tall; 
hydranths scarlet; up to an inch across tentacles when 
expanded; generally not found alive in summer 

Tubularia crocea 

Tubularia couthouyi 

Zanclea costata 

19. Colonies of very small, elongated hydranths with perisarc 

visible only at base; numerous capitate tentacles scattered 
over hydranths (fig. 6), with an apical whorl of 4-6 ten- 
tacles; color pink; height about 1 cm; free medusae budded 
from hydranths, below tentacles 

19. Irregularly branching colonies, with shorter bodied hydranths 
bearing about 16 capitate tentacles; free medusae produced 

from hydranth body close to tentacles Sarsia tubulosa 

20. Hydrothecae on pedicels or wineglass-like stems (fig. 3, 8) 21 

20. Hydrothecae lacking stems ("sessile", 'adnate "), closely 

appressed to stem or branch (fig. 5, 10) 31 



21. Hydrotheca, as expected, large enough to enclose hy- 

21. Hydrotheca aberrant in being very shallow saucer or 
funnel-shaped, usually marked by circlet of bright 
beadlike dots, not covering hydranths (fig. 19); stem 
fascicled; fanlike branching colony up to 10 inches 
high; female gonangia in rows on upper side of branch- 
es, each surmounted by a pair of hydranths (fig. 18) 


Halecium halecinum 

22. Hydrotheca tubular, and provided with a conical oper- 
culum of toothlike flaps (fig. 14); family Campanulin- 
idae, a difficult group of very small forms, of which 
the most common local form may be Calycella syringa , 
a tiny (1/g inch) species found on other hydroids, bry- 
ozoans, etc. 

22. Hydrotheca bell or wine-glass shaped, with inner dia- 
phragm or annular thickening at base of hydranth, but 
lacking operculum (closing device) : A very numerous 

Campanulariidae 2 3 

Cautionary note: The only basic way to define the genera of Campanulariidae 
is by their sexual stages. "It appears to be impossible to construct gener- 
ic characters for the Campanulariidae on the basis of the trophosome. The 
classification of the group is unnatural and unsatisfactory in the extreme, 
but this is not the place to attempt its rectification". (Nutting, 1901, 
p. 344). 

a. Produce free medusae having 4-8 tentacles at time of 

release Clytia (= Phialidium ) 

b. Produce tiny free medusae with 12-16 tentacles at time 

of release Qbelia 

c. Medusoids are very reduced and remain in gonotheca; 

hydrotheca with annular thickening at base Campanularia 

d. Medusoids are extruded from gonotheca in a sac (acro- 
cyst) but are not released; hydrotheca with thin dia- 
phragm at base of hydranth Gonothyraea (=Laomedea) 

23. Colonies consisting of unbranched or not regularly branched 
growths arising from a more or less extensive hydrorhizal 
system; hydrothecal margins toothed; gonangia borne on hy- 
drorhiza and ringed (annulated) , Japanese lantern fashion 
(figs. 3, 4); release 4-8 tentacled medusae 
Clytia 24 

23. Colonies regularly branching (at least in terminal twigs); 

gonangia not ringed 25 

24. Colony of unbranched (or occasionally branched) white 
growths arising from extensive rootstock; height about 
V4 inch; hydrothecal rim with ca. 16 rounded teeth; gon- 
angia deeply ringed (fig. 4) 

24. Colony white, profusely branched but without clear main 

stem; height about 1 inch; hydrothecal rim with 12-14 sharp 
teeth; gonangia irregularly ringed (fig. 3) 

Clytia johnstoni 

Clytia edwardsi 

;L6 Hydrozoa 

25. Gonangia liberate small medusae with 12-16 tentacles; 

several species , of which only 3 are in key Obelia 26 

25. Gonangia contain the reduced medusoids (sporosacs) 

which are never released; only 4 of the several species 

are in key Campanularia 28 

25. Gonangia extrude the reduced medusoids inside a sac 

(acrocyst) Gonothyraea (= Laomedea ) loveni 

Note: Of the campanulariid genera occurring at Woods Hole, Laomedea and Obel - 
ia can be distinguished by the possession of a thin diaphragm in the hydro- 
theca at the base of the hydranth. In the others ( Clytia and Campanularia ) 
this basal structure has the form of a ring shaped thickening. 

26. Main stem simple; hydrothecae smooth, shallow, tooth- 
less; gonangia with constricted opening, "bottle necked" 

(fig. 13) 27 

26. Main stem fascicled; hydrothecae deep, longitudinally 
ribbed, with 14-20 double-pointed teeth on rim; gonangia 
with opening not constricted; colony may exceed 30 inches 

Obelia bicuspidata 

27. Colony of stiff, short (1 inch or less) unbranched, zig- 
zag (geniculate) stems with short, stout internodes and 
hydranths set alternately, in one plane; gonangia about 

5 times length of hydrothecae Obelia geniculata 

27. Colony much branched, with long slender internodes; gon- 
angia about 3 times length of hydrothecae; height of col- 
ony 6-8 inches Obelia commissuralis 

28. Main stem with many branches, which rebranch; gonothecae 
with very small apertures; colony in absence of gonangia 
closely resembles Obelia commissuralis ; height about 6 

inches Campanularia amphora 

28. Stem bearing only pedicels, or a few branches 29 

29. Stem brownish and stout, with short internodes (4-5 times 
as long as thick, fig. 7); height about 1.5 inches; gono- 
thecae of tubular form with apertures not constricted 

Campanularia f lexuosa 

29. Main stems slender with longish internodes (7-8 times as 

long as thick, fig. 8) ; gonangia with specialized apertures 30 

30. Stem strongly zigzag (geniculate) ; height about 3/4 inch; 
gonangia irregularly wrinkled, bottle-necked 

Campanularia angulata 

30. Stem flexuose (weakly zigzag); height up to 1.5 inch; fe- 
male gonangia with a distinctive folded-over tip (fig. 9) ; 
male gonangia bluntly fusiform and irregularly bulging 

(fig. 8) Campanularia calceolifera 

31. Hydrothecae adnate on one side only of stem or branches 
(family Plumulariidae) ; the species commonly taken has a 
white, delicate feathery colony, 1-3 inches high; tiny 
special zooids (nematophores) form small spout-like ex- 
tensions set along branches between hydrothecae (fig. 5); 
gonangia curved, set among branches 

Schizotricha tenella 

31. Hydrothecae arranged on both sides of stem or branches 

(Sertulariidae) 32 

Hydrozoa 17 

32. Hydrothecae set as opposite pairs (genus Sertularia ) ; 
in the commonest local species, the members of each 
pair of hydrothecae not in contact (fig. 10); gono- 
theca bottle-necked, not annulated; dark brown, stiff 
growths, usually on seaweeds; 0.5-1.5 inches high 
Sertularia pumi la 

32. Hydrothecae set in alternating fashion 33 

33. Pinnately-branching brown colony, up to 12 inches tall; 
hydrothecae with tubular necks, smooth-rimmed 

Abietinaria abietina 

33. Colony dichotomously branching; a handsome bushy form 
("squirrel-tail hydroid") with silvery branches, up to 
12 inches tall; hydrothecal rims elevated into two op- 
posite teeth Thuiaria argentea 


In view of the large number of species alleged by Nutting and Fraser for this 
region, it would be premature to attempt a complete list. The experimentalist 
should be aware that species not on this list may be expected to turn up, that a 
given genus, e.g. Tubularia , might be represented in Supply Department collections 
by different species at different seasons, and that some species and even genera 
in certain families, notably Campanulariidae, are impossible to identify without 
sexually mature ("fruiting") material. 

Abietinaria abietina (Linnaeus, 1758) . Dredged. 

Acaulis primarius Stimpson, 1854. Rare, in dredgings north of Cape. 

Bougainvillia carolinensis (McCrady, 1858) . On rocks, pilings, algae; common. 

Bougainvillia superciliaris (L. Agassiz, 1849). Status unclear. According to Rus- 
sell (1953), hydroid is tiny, with ca. 6 filiform tentacles in a whorl, hy- 
dranths arising singly from stolon. Nutting (1901), following Agassiz, des- 
cribed hydroid as up to 2 inches high with 15-20 tentacles, but did not see it 
himself. If this species occurs here, it may occur chiefly as the medusa. 

Calycella syrinqa (Linnaeus, 1767). Common on other hydroids, bryozoans, algae, 
but inconspicuous because of small size. 

Clava leptostyla L. Agassiz, 1862. Most commonly in small clusters on Ascophyllum 
nodosum in low intertidal; of rather local, sporadic occurrence. 

Campanularia amphora (L. Agassiz, 1862). Common in shallow water. Easily confused 
with Obelia commis sura lis . 

Campanularia angulata (Hincks, 1861) . Reported on Zostera . 

Campanularia calceolifera H incks, 1871. Common on Mytilus , seaweeds, and pilings. 
The unique folded-over tip of female gonangium makes identification certain. 

Campanular ia f lexuosa (Alder, 1856) . On pilinqs, rocks, algae. Very common. 

Clytia edwardsi (Nutting, 1901). On pilings, etc. 

Clytia Johns ton i (Alder, 1857) . (= Clytia bicophora Nutting, 1901) . The medusae 

produced by the hydroid genus Clytia are known as Phialidium and Russell re- 
fers Clytia johnstoni in British waters to Phialidium hemisphericum Linnaeus. 
The disposition of the local Clytia johnstoni should await proper study. The 
hydroid is found locally on intertidal brown algae and on rocks. 
Note ; Clytia differs from Campanularia essentially only in the degree of de- 
velopment of its medusa. It is possible that future practice will be to in- 
clude Clytia within the genus Campanularia . 

Cordylophora lacustris Allman, 1844. An important experimental animal and the only 
typical colonial hydroid found in fresh water. On pilings, under floats, and 
in culverts in ponds and passages in areas of low salinity. 

C orynitis agassizi , see Z anclea costata . 

Corymorpha pendula L. Agassiz, 1862. Dredged quite rarely, from soft bottoms. 

18 Hydrozoa 

Ectopleura dumortieri (Van Beneden, 1844) . (= Ectopleura ochracea L. Agassiz, 1862) . 
This is generally seen as the medusa. The hydroid is a solitary form resem- 
bling Tubular ia , but producing free-swimming medusae. 

Eudendrium album Nutting, 1898. Small, inconspicuous; on rocks, piles, algae. 

Eudendrium carneum Clarke, 1882. A conspicuous species, with red hydranths and go- 
nophores. This and the next species are common laboratory examples of hydroids 
with greatly reduced medusoids. On piles and algae. 

Eudendrium ramosum Linnaeus, 1758. Also conspicuous. One of the commonest shallow 
water forms on rocks and piles, and in deeper water. 

Eudendrium tenue A. Agassiz, 1865. Shallow water, on piles. 

Gonothyraea (= Laomedea ) loveni (Allman, 1859) . On shells, stones, pilings, floats, 
in shallow water. 

Halecium halecinum (Linnaeus, 1767). Common in shallow water on shells, stones, 

Hydractinia echinata (Fleming, 1828) . Very common on shells of Littorina (see 

Crowell, 1945) and other snails occupied by hermit crabs, and also on rocks and 
piles. Seems quite variable, both in roughness of spines and in color, which 
varies from pure white to a rich salmon. 

Laomedea , see Gonothyraea . 

Obelia bicuspidata Clark, 1876. As with other campanulariids, generic identification 
of Obelia may be impossible without mature material. On Zostera , piles, and in 
deeper water. 

Obelia commissuralis McCrady, 1858. Widespread in shallow water on various sub- 
strates . 

Obelia geniculata (Linnaeus, 1758). Grows profusely on floats, piles, and Lamin- 
aria; on the latter the extensive stolon system is conspicuous. 

Pennaria tiarella (Ayres, 1854). A common and conspicuous species in clear shallow 
water under rock ledges, etc. A favorite laboratory animal. Bears reduced med- 
usae with 4 rudiments of tentacles. Medusae may be released, or may discharge 
gametes while still attached. In laboratory this is usually seen between 7 and 
9 P.M. 

Podocoryne carnea Sars, 1846. Generally collected on shells of Nassarius trivitta - 
tus used by hermit crabs (see Crowell, 1945), although not confined to this 
shell. Color pink to pure white. An important experimental animal in recent 
years. Since Podocoryne differs from Hydractinia mainly in the development of 
a free medusae, it is possible that future practice will be to include both in 
the genus Hydractinia. 

Sarsia tubulosa (Sars, 1835). This species is probably not to be found at Woods 
Hole in summer. Sumner ejt al^. cite records of its abundance in March and Ap- 
ril, with the medusae most common in April and May. The hydroid has not been 
included in the keys and lists of the Invertebrate Zoology Course for at least 
25 years. The free medusa is one of a difficult group and Russell includes 
Syncoryne mirabilis (L. Agassiz, 1849) as a synonym of Sarsia tubulosa . 

Schizotricha tenella (Verrill, 1874) . The delicate feathery form is easily recog- 
nized. Common on pilings. 

Sertularia pumila Linnaeus, 1758. Common on intertidal brown algae in protected 
waters . 

Stylactis hooperi Sigerfoos, 1899. Occurs on shells of living mud snails (Nas - 
sarius obsoletus ) . Since Stylactis , like Podocoryne , differs from Hydractinia 
mainly in the degree of development of its medusa, it is possible that future 
practice will be to include all three within the genus Hydractinia. 

Syncoryne mirabilis , see Sarsia tubulosa . 

T huiaria argentea (Linnaeus, 1758). Dredged, but usually dead and empty when taken 
in summer. 

Tubular ia : A difficult genus, but very important in experimental work. Specific 
identification is often inadequate, an unfortunate situation when results of 
different authors have to be compared. Experimentalists using Tubularia would 
do well to record date, water temperature, and place of collection of labora- 
tory material. Nutting lists five species; this key, three: 

Hydrozoa 19 

Tubularia couthouyi L. Agassiz, 1862. On sandy or stony bottom. This large and 
beautiful species has generally died off by summer. 

Tubularia crocea (L. Agassiz, 1862). Subtidal, on pilings, sometimes in brackish 
water. The same species is thought to occur in the Oakland Estuary (San Fran- 
cisco Bay) . This is the species most used experimentally. 

Tubularia larynx Ellis and Solander, 1786. On rocks, piles, and algae. 

Turritopsis nutricula (McCrady, 1856). This is generally seen as the medusa. The 
hydroids are small and inconspicuous. 

Zanclea costata Gegenbaur, 1856. The work of Russell makes it seem probable that 
the hydroid " Corynitis agassizi " and the medusa Gemmaria gemmosa as recorded 
from Woods Hole should be referred to Zanclea costata . This is found abun- 
dantly on the red bryozoan nodules obtained by dredging. 


Numerous small medusae occur seasonally in the plankton of Woods Hole and ad- 
jacent waters, but their identification is a matter of considerable difficulty. 
One may identify a good many simply by reference to the illustrations in Hargitt 
(1905), but there remain problems of synonymy because life histories of medusae 
were not well worked out at that time, and indeed are still very incompletely 
known. Reference should be made to Mayer's "Medusae of the World" (1910). A use- 
ful guide to the generic identification is the "Pictorial key to species of British 
Medusae", pp. 42-45 in the excellent volume by Russell (1953). In the present man- 
ual, no key has been attempted. 

1. Order HYDROIDA (includes most of the local hydromedusans) : 

a. Suborder ANTHOMEDUSAE : These are the medusae of the gymnoblastic or athe- 

cate hydroids, and are characterized by a deep, bell shaped form, lacking 
statocysts, and with the gonads born on the manubrium. The medusae of 
Bouqainvillia and Podocoryne are typical examples (Plate 2, figs. 1 and 15). 

b. Suborder LEPTOMEDUSAE : These are the medusae of such of the calyptoblas- 

tic or thecate hydroids as produce medusae (most do not) . They are of flat- 
tened form with gonads born on the radial canals, and ectodermal statocysts. 
Obelia and Phialidium (= Clytia ) are examples. 

c. Suborder LIMNOMEDUSAE : This group is not recognized as such by Hyman (1940) 

and is given ordinal rank by Russell (1953) . It includes the well known 
Craspedacusta sowerbii , of sporadic occurrence in fresh waters; its polyp 
stage is the minute, tentacle-less " Microhydra " . Another very famous rep- 
resentative, and the subject of much experimental work at Woods Hole, is 
Gonionemus vertens A. Agassiz, 1862. This was once very abundant in the Eel 
Pond, but it became very scarce in the Woods Hole region with the dying off 
of the eel grass ( Zostera ) about 1930, and, despite the general return of 
the grass, is now of rather sporadic and unpredictable occurrence. In some 
summers thousands may be seen in shallow, protected bays on Martha's Vine- 
yard; in other years very few may be taken. Most of the Woods Hole papers 
on this species refer to it as Gonionemus murbachi , which Kramp (1959) re- 
gards as a synonym of G. vertens . 

2. Order TRACHYLINA (Suborders TRACHYMEDUSAE and NARCOMEDUSAE) : Trachyline medusae 
are generally oceanic and seldom taken in the inshore waters near Woods Hole. 
They lack a true hydroid stage, or may have a parasitic larval development; 
adults may have a distinctive scalloped bell margin, or tentacles inserted above 
the margin. Russell's pictorial key may be helpful if members of this group are 

The last two yroups to be mentioned are commonly called "siphonophores " . 
These (s ensu lato ) are floating or swimming colonies containing both polypoid 
and medusoid individuals. The recent trend in classification is to split this 

2 Hydromedusae 

group into two orders : 

3. Order CHONDROPHORA, allied to the tubularian hydroids, and represented in this 
region only by rare examples of the "purple sailor", Velella velella (Linnaeus), 
which has an oval float with diagonal vertical sail, and the somewhat similar 
Porpita porpita (Linnaeus) , which lacks a sail. 

4. Order SIPHONOPHORA proper, represented locally by the "Portuguese man-of-war", 
Physalia physalia (Linnaeus) , which is well known and easily recognized by its 
large purple and rose irridescent float. Physalia may be quite common in Vine- 
yard Sound in certain summers, especially after long periods of southeast storms. 
However, experimentalists are advised that in occasional summers it is virtually 
not to be found. The sting is severe and, although they may be picked up by the 
float, they should be handled with caution and avoided by swimmers. Other si- 
phonophores are of rare or occasional occurrence. 


Crowell, S., 1945. A comparison of the shells utilized by Hydractinia and Podo- 

coryne . Ecology, 26 :207. 
Fraser, C. M., 1944. Hydroids of the Atlantic Coast of North America > Univ. of 

Toronto Press, 451 pp., 94 pi. 
Hargitt, C. W. , 1905. The Medusae of the Woods Hole region. Bull. U. S. Bur. 

Fish., 1904, 24: 21-79, pi. 1-7. 
Hyman, L. H., 1940. The Invertebrates, vol. I, Protozoa through Ctenophora, 

726 pp., McGraw-Hill. 
Kramp, P. L., 1959. The Hydromedusae of the Atlantic Ocean and adjacent waters. 

Dana Rept. 46_: 1-283, Pi. I -II. 
Mayer, A. G., 1901. The Medusae of the World. Vols. I and II, The Hydromedusae. 

Carnegie Inst, of Wash. Publ. 109 . 
Nutting, C. C, 1899. The hydroids of the Woods Hole region. Bull. U. S. Bur. 

Fish., 1899, _19: 325-386. 
Russell, F. S., 1953. The Medusae of the British Isles, Cambridge Univ. Press, 

530 pp., 35 pi. 
Totton, A. K., and G. 0. Mackie, 1960. Studies on Physalia physalis (L.). Part I., 

Natural history and morphology. Part II. Behavior and histology. Discovery 

Repts, 30: 301-408. 

Chapter III 21 


The commonly encountered members of the Class Scyphozoa are well known to the 
general public as " jellyf ishes" , and are often feared unreasonably. And it is fair 
to advise that any large medusa should be handled with due respect, for its sting- 
ing powers despite exaggerations in popular literature, may be formidable (cf. 
Hedgpeth, 1945). 

Scyphozoan jellyfish are commonly large and conspicuous, with scalloped bell- 
margins and usually with long tentacles and/or mouth lobes. The attached or poly- 
poid stages are absent or inconspicuous; these "scyphistoma" stages (Plate 3, fig. 
3) may occasionally be found on eelgrass, rocks, or timbers, or can be reared from 
larvae shed by medusae in the laboratory. 

Members of the order Stauromedusae are easily distinguishable from other mem- 
bers of the Class in that they are sessile throughout their life cycle except for 
a brief period during development when a creeping, vermiform, nonciliated planula 
larva is produced. Two families have been designated, the Eleutherocarpidae, rep- 
resented locally by Haliclystus , and the Cleistocarpidae, represented by Cr-atpro- 
lophu s. The latter group possesses "claustra", or membranes, which divide the gas- 
tric pockets (see figs. 162B, p. 502, and 165J, p. 510 in_ Hyman, The Invertebrates , 
Vol. I, for the distinction between the two families). Two stauromedusans are found 
locally early in the season (to the end of June) attached to Fucus at Nobska Point. 

The synonymy of Scyphozoa is extensive, and most of the names used by Hargitt 
and by Mayer have been modified. In the keys below, the names in Kramp's synop- 
sis (1961) have been used. For the key to attached scyphozoans we are indebted 
to Drs. G. F. Gwilliam and Kay Werner Petersen; the key to scyphomedusae has been 
derived from Mayer (1910). Figure references are to Plate 3. 


1. Small (under 10 mm) soft individuals connected by stolons 
(fig. 3); up to 16 or more very long filiform tentacles; 
gastric cavity with 4 small radial septa 
"Scyphistoma" stages of various scyphozoan medusae 

1. Larger (up to 30 mm) ; broadly bell shaped single indiv- 

uals; 8 marginal clusters of short knobbed tentacles . . Order STAUROMEDUSAE 2 

2. Stalk comparable in length to depth of bell; with conspic- 

uous marginal anchors (fig. 2) between tentacle-bearing 

arms Haliclystus auricula (Rathke, 1806) 

2. Stalk much shorter than depth of bell; no marginal anchors 

(fig. 1) Craterolophus convolvulus (Johnston, 1835) 

Note: If the stauromedusan was collected north of Cape Cod, consult 
the paper of Berrill (1961) before concluding an identification. 


1. Bell flat, translucent grayish, with 4 horeshoe shaped 
gonads; marginal tentacles very short and numerous 
Aurelia aurita (Linnaeus, 1758) 

1. Bell deep and usually colored; marginal tentacles long 2 

2. Marginal tentacles long and numerous, arranged in 8 

clusters, each of several rows Cyanea capillata (Linnaeus, 1758) 

2. Marginal tentacles single 3 


Plate 3 


Fig. 1. Craterolophus convolulus , from a preserved specimen; 
bar is 1 cm. 

2. Haliclystus auricula , from a preserved specimen; bar 
is 1 cm. 

3. Scyphistoma stage of Aurelia aurita , grown in labora- 
tory and sketched from life by Dr. Louise Bush. Scale 
is about 1 millimeter. 

4. Astrangia danae , a small section of living colony, 
sketched from life by Bruce Shearer. Scale is 5 mm. 

5. Haloclava producta , after Hargitt; this and following 
3 figures redrawn by Mrs. Emily Reid. Scale bars of 
figures 5-8 approximately 1 cm long. 

6. Sagartia modesta , after Hargitt. 

7. Nematos Leila vectensis , after Crowell. 

8. Edwardsia elegans , after Hargitt. 

Plate 3 

I cm 


24 Scyphomedusae 

3. Eight marginal tentacles; 16 marginal lappets; bell 
with numerous nematocyst bearing warts 

Pelagia noctiluca (Forskal, 1775) 

3. Up to 40 marginal tentacles; about 48 marginal lappets 

Chrysaora quinquecirrha (Desor, 1848) 


Berrill, M., 1962. The biology of three New England Stauroraedusae, with a descrip- 
tion of a new species. Canad. J. Zool., 4_1: 1249-1262. 

Hargitt, C. W„, 1905. The medusae of the Woods Hole Region. Bull. U. S. Bur. Fish. 
1904, 24: 21-79. 

Hedgpeth, J. W. , 1945. Re-examination of the Adventure of the Lion's Mane. Sci. 
Monthly, 60: 227-232. 

Kramp, P. L., 1961. Synopsis of the medusae of the world. J. Mar. Biol. Assoc. 
U. K.j 40: 7-469. 

Mayer, A. G., 1910. Medusae of the World. Vol III, The Scyphomedusae. Carnegie 
Inst, of Washington, Publ. no. 109. 

Chapter IV 25 


By Cadet Hand 

This, the largest class of the Cnidaria, has but a modest representation at 
Woods Hole, chiefly in the group of sea anemones. 


Members of this subclass are at once identifiable by the possession of 8 pin- 
nate tentacles. The group contains several orders, mostly of warmer waters, such 
as sea pens, sea fans, gorgonians, "organ-pipe coral", the sea pansy ( Renilla ) , 
well known in experimental work, and the precious red coral of jewelry. In the 
Woods Hole region, dredging off Martha's Vineyard and in the Sound may yield two 
species . 

1. Order ALCYONACEA ("soft corals"): Alcyonium carneum Agassiz. 1850. Grows 
as fleshy, finger-like lobes attached to rocks and shells at depths of 20-60 me- 
ters; the color varies from bright salmon to pink or flesh-colored. The tissue 
contains calcareous spicules, but there is no firm skeleton. At collection the 
polyps are retracted and the general appearance is describable by the common 
name of "dead men's fingers". In the relaxed condition the delicate polyps re- 
veal the 8 pinnate tentacles characteristic of the subclass. It survives only 
moderately well under laboratory aquarium conditions. 

2. Order PENNATULACEA : Pennatula aculeata Danielson and Koren, 1858. This 
"sea pen" has the form of a feather, with a bulbous base on which it stands in 

soft muddy bottoms. The local species, which is taken only occasionally, is 
10-25 cm in height and of a purplish-red color. 

B. Subclass ZOANTHARIA: 

Here are placed the great assemblages of stony corals (Madreporaria) , sea 
anemones (Actinaria) , and some other orders, examples of which are rare or lacking 
in the Woods Hole region. 

1. Order MADREPORARIA ("true" or stony corals): Astrangia danae Agassiz , 1847 
is the sole local representative. It is a typical coral in form (Plate 3, fig. 
4) forming irregular encrustations or low branching growths almost everywhere on 
rocks, shells and pilings from low-water mark to 30-40 meters. It is hardy, and 
lives well in laboratory sea-water aquaria. 

2. Order CERIANTHARIA: Cerianthus americanus Verrill. 1866. A southern form, 
this has been taken only very rarely at Woods Hole. It has the form of an elon- 
gated (up to 20 cm) burrowing sea anemone with 2 whorls of tentacles, living in 
a distinct mucoid tube. McMurrich (1890, J. Morph. , 4: 131-150) gives a good 
colored illustration. See Field (1949) for additional information. 

3. Order ZOANTHIDEA: Members of this order resemble small sea anemones but 
are generally colonial. Epizoanthus americanus (Verrill, 1864) is barely in- 
cludable in the Woods Hole region, being common only at depths of 50 or more 
meters, off Martha's Vineyard and Nantucket. It generally occurs on shells oc- 
cupied by hermit crabs, and also on rocks. 

4. Order ACTINIARIA (sea anemones) : These are familiar animals, generally 
recognizable as such. Several local species are readily identifiable from the 
key My form, color, and habitat, but critical identification of the rest is made 
difficult by the fact that classification is based upon the assemblage of nema- 
tocyst types ("cnidome") and upon internal anatomy such as mesenterial arrange- 
ment, musculature, and other details requiring microscopical sections. The stu- 
dent who wishes to go beyond the following key would do well to consult the works 
of Stephenson (1928, 1935) and Carlgren (1949). 

26 Anthozoa 

(Figure references are to Plate 3) 

1. Burrowing or buried in sand, gravel, or mud with only 

the tentacles exposed; body elongate, even worm-like 2 

1. Attached on hard surfaces with most of body well ex- 
posed 5 

2. Upper part of body with 20 rows of papillae (fig. 5); 
tentacles 20 in number, blunt and tending to be swollen 
at tips; up to 10-15 cm body length 

Haloclava producta 

2. Tentacles pointed and body without papillae 3 

3. Acontia* present; flattened base attached to pebbles or 

shells (fig. 6) Sagartia modes ta 

3. No acontia; aboral end rounded or pointed; no flattened 

base; normally 16 tentacles 4 

4. Small, not over 2 cm long; transparent when extended and 
without adherent sand (fig. 7) 

Nematostella vectensis 

4. With middle portion of body covered by a rough brown 
cuticle to which sand adheres; 8 longitudinal grooves (fig. 

8) ; body up to 3.5 cm long Edwardsia elegans 

5. Column green to grayish black, with or without orange, 

yellow, or white lines Haliplanella luciae 

5. Coiutin white to salmon or brown, not green or black, 

sometimes mottled with brown 6 

6. Large individuals with lobed or frilled tentacular 
crown. Tentacles very numerous and short. In exten- 
sion the column is usually less than twice as long as 

its diameter Metridium senile 

6. Column elongated when extended and usually 3 or more 
times longer than its diameter. Tentacles elongate. 
Color white to pinkish, sometimes showing a greenish 
tinge in the tentacles and upper column. Some speci- 
mens may show one or more larger and more opaque inner 
tentacles known as "catch tentacles", of uncertain func- 
tion Diadumene leucolena 

*) Acontia are ciliated, thread-like structures which arise from the free edges 
of the mesenteries and project into the gastric cavity. When acontiate anemones 
are roughly handled the acontia may be extruded through the mouth or through 
special pores (cinclides) in the body wall. 


With some exceptions the sea anemones of Woods Hole are not well known, and a 
restudy of local species is needed. The original descriptions, based largely on 
form and color, are inadequate by modern standards. 

Anemonia sargassensis Hargitt, 1908. Not in key; an occasional visitor upon drift- 
ing Sargassum . 

Bicid ium parasitica , see Pea chia parasitica . 

Diadumene l eucolena (Verrill, 1866). Listed as Cylista in Hargitt (1914). Most 
commonly found under stones on rocky beaches. See Hand (1955) for synonymy. 

Anthozoa 27 

Edwardsia elegans Verrill, 1869. Reported burrowing in intertidal sand flats (Har- 
gitt) ; apparently not common. 

Edwardsia leidyi V errill, 1898. This name applies to a larval stage found parasit- 
ic in the ctenophore Mnemiopsis leidyi . The adult has not been determined. 
Length 2-3 cm, color pink. 

Eloactis producta , see Haloclava producta . 

Halcampa duodecimcirrata (Sars, 1851) . One specimen, reported by Verrill dredged 
in mud off Gay Head, had 10 tentacles. Species was described as having 12 ten- 
tacles, from Maine specimens. On present evidence, rare, not in key. 

Halcampa farinacea , see Ha lcampa duodecimcirrata . 

Haliplanella luciae (Verrill, 1898) . This common and well known species has a very 
discouraging synonymy, having been assigned to the genera Sagartia , Chrysoela , 
Diadumene , Aipaisiomorpha , and Haliplanella , without there being much question 
of its specific name luciae (after Verrill's daughter Lucy) since Verrill's des- 
cription. H. luciae is thought to be of Japanese origin; it appeared near New 
Haven in 1892 and has since become one of the commonest anemones of New England; 
the first British record was in 1896, and it has also become widespread on the 
California coast. The color is variable: dark brown, olive or green, with or 
without lines of orange, yellow, or white. The orange lines when present, are 
diagnostic, but the unlined individuals are more difficult to identify. Very 
common, on pilings, among mussels, in salt marshes, often in brackish water. 
See Hand (1955) for synonymy. 

Haloclava producta (Stimpson, 1856). A burrowing form on intertidal sand flats. 

Metridium dianthus , fimbriatum , m arginatum , see Metridium senile . 

Metridium senile (Linnaeus) . In systematic work this is referred to as M. senile 
senile , while the Pacific coast subspecies is M. senile fimbriatum . The spe- 
cies at Woods Hole has also been variously called M. dianthus , M. marginatum , 
and M. fimbriatum , a not unusual situation when a very widespread and variable 
form is concerned. See Hand (1955) for synonymy. 

Nematostella vectensis Stephenson, 1935 (=N. pellucida Crowell, 1946) . A small deli- 
cate form found in the Mill Pond, Woods Hole, where salinity may fluctuate great- 
ly. Up to 20 mm long and 2-4 mm in diameter when extended. Lives in soft mud 
with disc and tentacles exposed. Crowell' s account of anatomy is excellent. 
See Hand (1957) for synonymy. 

Peachia parasitica (L. Agassiz, 1859) . Young stage reported parasitic in the jelly- 
fish Cyanea . Adults not yet recorded south of Eastport, Maine. Not in key. 

Sagartia leucolena , see Diadumene leucolena . 

Sagartia luciae, see Haliplanella luciae . 

Sagartia modesta Verrill, 1866. There is some doubt as to the proper generic name 
for this species; see Carlgren (1950) for discussion of this problem. 


Carlgren, 0., 1949. A survey of the Ptychodactaria , Corallimorpharia . and Actin - 

iaria . K. Svenska Veten. Handl. 4th ser., _1(1) : 1-121. 
Carlgren, 0., 1950. A revision of some Actiniaria described by A. E. Verrill. J. 

Wash. Acad. Sci., 40(1): 22-28. 
Crowell, S., 1946. A new sea anemone from Woods Hole, Massachusetts. J. Wash. 

Acad. Sci., 3j6: 57-60. (good account of Nematostella ) . 
Field, L. R., 1949. Sea anemones and corals of Beaufort, North Carolina. Dake 

Univ. Mar. Sta. Bull. 5, 39 pp, 10 pi. (useful for Cerianthus but otherwise 

not much help in the Woods Hole region) . 
Hand, C, 1955. The sea anemones of central California. Part 111. The ^contia- 

rian anemones. Wasmann J. Biol., L3: 189-251. (synonymies and redescriptions 

of several Woods Hole species) . 
Hand, C, 1957. Another seaanemone from California. J. Wash. Acad. Sci., 47 : 

411-14. (Nematostel la) . 

28 Anthozoa 

Hargitt, C. W., 1914. The Anthozoa of the Woods Hole region. Bull. U. S. Bur. 

Fish., 32 (for 1912): 223-54. PI. 41-44. 
Stephenson, T. A, 1928. The British Sea Anemones. Vol. I, 148 pp. (Ray Soc. Vol. 

113) London. 
Stephenson, T. A., 1935. The British Sea Anemones. Vol. II, 426 pp. (Ray Soc. 

Vol. 121) . London. 

Admonitory note : Generic name changes may occasionally lead the experimentalist 
using this manual to give way to Frustration and Despair. Let him remember, how- 
ever, that systematists are professional gentlemen who are doing the best they 
can, supported by such declarations as the following: 




VOLUME I. Part 4. Pp. 23-30. 



On the need for avoiding intemperate language 

in discussions on zoological nomenclature 

DECLARATION. — In the opinion of the Commission the tendency to enter into pub- 
lic polemics over matters which educated and refined professional gentlemen might 
so easily settle in refined and diplomatic correspondence is distinctly unfavor- 
able to a settlement of the nomenclatorial cases for which a solution is sought. 
It may be assumed that the vast majority of zoologists agree with the Commission 
in desiring results rather than polemics, and the Commission ventures to suggest 
that results may be obtained more easily by the utmost consideration for the usual 
rules of courtesy when discussing the views of others. 

The dangers attending the use of sarcasm and intemperate language in discus- 
sions on zoological nomenclature were specially considered by the International 
Commission on Zoological Nomenclature at their Session held at Monaco in March 
1913 during the Ninth International Congress of Zoology. The Commission consid- 
ered that this guestion was sufficiently pressing to reguire special treatment in 
their report to the Congress. In framing that report the Commission accordingly 
devoted paragraphs (68) and (69) to this subject. 

2. Paragraph (68) of that report reads as follows: — 

(68) Intemperate Language . --Whether or not it be an actual fact, appearances 
to that effect exist that if one author changes or corrects the names used by an- 
other writer, the latter seems inclined to take the change as a personal offense. 
The explanation of this fact (or appearance, as the case may be) is not entirely 
clear. If one person corrects the grammar of another, this action seems to be in- 
terpreted as a criticism upon the good breeding or education of the latter person. 
Nomenclature has been called "the grammar of science" and possibly there is some 
inborn feeling that changes in nomenclature involve a reflection upon one's educa- 
tion, culture and breeding. Too freguently there follows a discussion in which 
one or the other author so far departs from the paths of diplomatic discussion, 
that he seems to give more of less foundation to the view that there is something 
in his culture subject to criticism. It is with distinct regret that the Com- 
mission notices the tendency to sarcasm and intemperate language so noticeable 
in discussions which should be not only of the most friendly nature, especially 
since a thorough mutual understanding is so valuable to an agreement, but which 
are complicated and rendered more difficult of results by every little departure 
from those methods adopted by professional gentlemen. 

Chapter V 



The "comb jellies", although classified as coelenterates, are now generally regarded 
as a phylum distinct from the proper coelenterates or Cnidaria. Ctenophores possess 
biradial symmetry, an aboral sensory area, and 8 meridional rows of fused cilia (paddle- 
plates) ; they lack the circumoral tentacles and nematocysts characteristic of the Cnidar- 
ia, and show none of the polymorphism and attached "polypoid" stages so characteristic 
of the latter phylum. They are essentially marine, although some are common in brack- 
ish waters as well. 

1. Possessing a pair of tentacles, each with small side branches 
1. Without tentacles; body in the form of a flattened sac with 


wide mouth 

Class NUDA, Order Beroida 4 

2. Body of simple outline, without oral lobes; tentacles long but 

retractile into sheaths; gastric branches end blindly Order Cydippida 3 

2. Body with 2 large oral lobes, and 4 smaller lobes (auricles), 

body somewhat triangular in outline; tentacles inconspicuous, 

without sheaths; gastric branches fuse to form loops in oral 

lobes; Order Lobata, Mnemiopsis leidyi 

Body firm, egg shaped to nearly spherical; up to 20 mm long; 
not flattened; tentacles long, with many side branches 

3. Body of oval outline but compressed, resembling a flattened 
Pleurobrachia; rare 

Pleurobrachia pileus 

Mertensia ovum 

4. The side branches of the 8 meridional gastric canals beneath 
the comb rows do not anastomose; generally north of Cape Cod 

4. Side branches of the 8 meridional gastric canals anastomose 

Beroe cucumis 

Beroe ovata 


Beroe cucumis Fabricius, 1780. A common species on the northern New England coast; rare 
or occasional south of Cape Cod. 

Beroe ovata Chamisso and Eysenhardt, 1821. A southern species, irregular in occurrence 
north of Delaware Bay. 

Mertensia ovum (Fabricius, 1780). An arctic species, the young of which occur occasion- 
ally as far south as New Jersey. 

Mnemiopsis leidyi A. Agassiz , 1865. This is the commonest ctenophore of Woods Hole 

waters, especially in late summer and fall. Brilliantly luminescent when disturbed. 
Often found to contain wormlike pink, immature stages of the sea anemone Edwardsia sp. 

Pleurobrachia pileus (Fabricius, 1780). An arctic species, found atWjocsHole in winter 
and spring; usually not seen in July and August. 


Hyman, L. H. , Chapter VIII, Ctenophora, in_ "The Invertebrates", Vol. I, Protozoa through 

Ctenophora, 1940. 
Mayer, A. G. , 1911. Ctenophores of the Atlantic Coast of North America. Carnegie Inst. 

of Washington, Publ. no. 162 : 1-58, 17 pi. 

30 Chapter VI 


Class Turbellaria 

The Platyhelminthes comprise three classes of which two, the Trematoda (flukes) 
and Cestoda (tapeworms), are parasitic and although well represented in the Woods 
Hole region are not treated in this manual. Of the mostly free living class Tur- 
bellaria, a few species of the orders Tricladida and Polycladida are well known to 
investigators at the MBL, but there are a great many small forms belonging to the 
orders Aceola, Rhabdocoela, and Alloeocoela which are not well known, and which 
merit much further investigation. The work of Dr. Louise Bush in preparing the pre- 
liminary key, check list, and illustrations of these smaller forms is gratefully 


by Dr. Louise Bush 

Turbellarians may be collected in several ways. Many of the larger forms will 
come to the top of the water or crawl to the surface of mud or debris which has been 
allowed to stand in containers in the laboratory for several hours or days. This is 
the time honored method and works well for large marine species and for many fresh 
water forms, but the smaller marine species do not survive crowding or standing in 
containers with other plants and animals for more than a few hours; to obtain these, 
materials must be examined as soon after collection as possible. Stones, algae, de- 
bris, and especially hydroids and other materials from pile scrapings should be washed 
or stirred up thoroughly in sea water and the water poured off and examined under a 
dissecting binocular. Turbellaria may be found in such washings swimming about, mov- 
ing beneath the surface film, or crawling about on the bottom or sides of the con- 
tainer. Hand sorting of debris under a dissecting microscope may also be resorted 
to, but does not seem to be more effective than washing and is much more time con- 

Examination of the animals should first be made as they swim or crawl about. A 
familiar species may often be recognized by color, shape, and type of movement, but 
for exact determination, careful study of compressed specimens, of permanent mounts, 
and often of serial sections is necessary. The general student may make a tentative 
determination and learn something of the structure of these animals by mounting them 
on a slide so that they are slightly flattened by the coverslip. This may be accom- 
plished by judicious control of the amount of water on the slide or by ringing the 
coverslip with vaseline and compressing the material while observing it under the 

The following key uses characters which may be easily seen, and should enable 
the student to identify some of the commoner species found at Woods Hole, but in 
cases where examination of permanent mounts or sections is mandatory for specific 
identification the key will lead only to some of the higher categories. Figure 
references are to Plate 4. 


by Dr. Louise Bush 

1. Large: over 5 mm in length. Body flattened and leaflike 
and usually showing the pattern produced by the ramifica- 
tions of the gut (other color patterns may also be present) : 

Order TRICLADIDA, see annotated list on page 32. 

Order POLYCLADIDA, see keys to species on page 36. 

1. Small: under 5 mm and often from 1-2 mm in length. Body 
sh-'-'pe varies from oval and flattened to cylindrical to 
almost globular or egg shaped 2 

Turbellaria 31 

2. Without a clearly defined gut; food lies in center of 
body more or less irregularly surrounded by parenchyma 
cells. Statocyst present but eyes absent 
Order ACOELA 3 

2. With a true gut which can be distinguished by its clear- 
ly defined outer wall. Eyes may be lacking or one or 
two pairs present. Statocysts only occasionally present 


3. Posterior end of body with indentation in which are 
found 1-5 small tail-like appendages (fig. la, b, c) . 
Color reddish-orange to orange 

Polychoerus caudatus 

3. No tail-like appendage present 4 

4. Yellowish area at anterior end of body with dark red- 
dish-purple to black pigment scattered in the middle 
of body. Under higher magnification this pigment is 
seen to be in violet to purple individual cells (fig. 2) 

Aphanostoma diversicolor 

4. Color otherwise 5 

5. Penis doubled and appearing as two distinct structures 
side by side at the very posterior end of body (fig. 3) . 
There may be more than one species of this genus in our 
area but at least one has elongated hairs or spines over 
the surface of the body and a yellow color from pigmented 
cells in the parenchyma 

Childia groenlandica 

5. Penis single and this, with accessory structures, may be 

seen in posterior third of the body 6 

6. Color orange to dark orange or reddish -brown, the pig- 
ment occurring in irregularly shaped little nodules 
over the surface of the body. Found creeping on the 
surface of mud below low tide (fig. 4) 

Anaperus gardineri 

6. Color pale tan to transparent. Sperm masses and eggs 
easily seen scattered at posterior end of body: several 
species of small acoels, mostly belonging to the family 


7. Asexual repioduction evident as transverse partitions 
formed or partly formed at intervals along the rather 
transparent body (fig. 5). Sexual reproduction rare. 

Eyes lacking Micros tomum davenporti 

7. Sexual reproduction only; no transverse divisions 8 

8. Statocyst present. Body elongated with a squarish post- 
erior end containing adhesive papillae, the whole being 
used for attachment (fig. 6). See check list for com- 
ment on this form Monocelis sp. 

8. Statocyst absent 9 

9. Body chalky white, with or without pigmented areas. 

Form "chunky", almost circular in cross section 10 

9. Body more or less transparent or otherwise colored but 
not chalky white. Form usually somewhat flattened 
or more or less elongated 11 












One pair of eyes with a network of black pigment between 
and around them so that the animal appears to have a 
distinct black spot on the anterior end. Pharynx just 
behind eyes. Penis without cuticular stylet at poster- 
ior end of body (fig. 7) Plagiostomum sp. 

Two pairs of eyes. A ciliated groove extends across 
body just behind the eyes. Pharynx in posterior half 

of body (fig. 8) Monoophorum sp. 

One pair of eyes without associated black pigment. 
Ciliated groove lacking 
young of Plagiostomum sp. or other alloeocoels 

Proboscis at anterior end of body, protruded through a 
pore at the apex in catching prey. Pharynx is separate 

and located near the middle of the body 

Tongue-like process lying in front of the eyes in the 
cavity with the pharynx, may be protruded through the 
subapical mouth (fig. 9) 


Woodsholia lilliei 

No protrusible proboscis or tongue-like process present 13 

A straight penis stylet appearing like a conspicuous 
needle at posterior end of body (fig. 10) 

Gyratris sp. 

Penis stylet otherwise; may be curved or coiled 

other species of the KALYPTORHYNCHIA 

Pharynx simple, that is, not conspicuously muscular. 
Mouth appears as a more or less oval opening just back 
of eyes. Penis stylet curved, needle-like, at poster- 
ior end of body. Adhesive papillae on rounded poster- 
ior end of body (fig. 11) 

Macrostomum sp. 

Pharynx doliiform, that is, appears as a bulbous mus- 
cular organ just posterior to the eyes (fig. 12) 

one of the species of DALYELLIOIDA 

Pharynx rosulate, that is, round and closing as if by a 
purse string. Mostly fresh water species 

various species of the TYPHLOPLANOIDA 

by Dr. Louise Bush 
Order Acoela 

Anaperus gardineri Graff, 1911. This species has been taken since 1953 for use in 
the Invertebrate Zoology Course from mud brought in from below low tide mark 
in Great Harbor. Many of these worms come to the surface after the mud has 
stood for two or three days in the laboratory. They do not seem to be present 
in the Eel Pond. 

Aphanostoma diversicolor Oersted, 1845. This species occurs on both sides of the 

North Atlantic, and is common at Woods Hole on algae. It is easily recognized 
by its coloration and by the more pointed ends of the body as compared to other 
common acoels. 

Childia groenlandica (Levinsen, 1879). Hyman (1959) considers the Childia spinosa 
described by Graff (1911) from Woods Hole to be a synonym of the widespread 
C_. groenlandica . 

P olychoerus caudatus Mark, 1892. Formerly common in the Woods Hole region and reg- 
ularly taken both in dredgings and on Ulva from pilings and rocks. Recently 

Turbellaria 33 

it has not been reported. Its use in embryological studies is described in Cos- 
tello et al. (1957) . 

Order Rhabdocoela 
Suborder Opisthandropora 

Macros tomum spp. Species of this genus in the Woods Hole fauna are probably 

most easily recognized by the characteristic curved penis stylet and the simple 
pharynx located at the anterior end of the gut close behind the eyes. Specimens 
common on Fucus at Nobska Point have adhesive papillae at the caudal end and 
conspicuous hairlike setae scattered among their cilia. 

Microstomum davenporti Graff, 1911. These animals have a characteristic appearance 
and are easily recognized, since they are one of the few marine flatworms (the 
only ones seen at Woods Hole) which regularly reproduce by transverse fission. 
The cylindrical transparent body with the gut as a straight tube inside shows 
clearly the development of transverse walls and new pharynges as division pro- 
ceeds. When handled, those individuals in which division is nearly completed 
usually break in two, so that one often gets only very short specimens on a 

Suborder Lecithophora 

Section Dalyellioida: Members of the family Dalyelliidae itself are mostly 
fresh water forms, but the section includes a number of marine genera and species, 
many of which are in the family Umagillidae and are endocommensal with marine in- 
vertebrates, especially echinoderms. The group is poorly known at Woods Hole. 

Section Kalyptorhynchia : Includes Gyratrix sp. , characterized by having a pro- 
boscis in a separate pocket opening anteriorly (fig. 10), and easily seen in the 
living animal. 

Section Typhloplanoida: This group probably includes Woodsholia lilliei Graff, 
1911, of which a few specimens are believed to have been taken in 1963. These have 
a tongue-like process, difficult to see, in the pharynx. The penis sheath is char- 
acteristic, and best seen in whole-mounts. 

Order Alloeocoela 

Suborder Cumulata: Species of this group seen at Woods Hole are thick and more 
or less cylindrical in body form, chalky white in color, sometimes with black or 
brown spots, stripes, bands, or reticulations. 

Monoophorum sp. Common on Ulva ; resembles a chalky white lump until disturbed, when 

it swims slowly away as a short cylindrical animal with two pairs of eyes and a 

stumpy tail. 
Plagiostomum sp. Longer than Monoophorum , and with only one pair of eyes, which are 

so surrounded with black pigment that the head appears to have a single large 

dorsal spot. 

Suborder Seriata 

Monocelis sp. In 1911, Graff described from the Eel Pond amonoce lid which he placed 
in a new genus, Myrmeciplana , distinguished from Monocelis by having sensory 
hairs born on sensory papillae on the rostral end of the body; his species was 
reported as being eyeless. Animals collected from the Eel Pond in 1963 resemble 
Myrmeciplana , but have a pair of eyes in most cases, and also have the anterior 


Figure 1. 

lb and lc . 

Plate 4 

TURBELLARIA (except for po lye lads) 

Figures lb, lc, and 15 are redrawn from Verrill 
(1892); rest redrawn from sources as cited or from 
life by Dr. Louise Bush. Scale bars on all fig- 
ures equal 1 mm. 

Polychoerus caudatus , with one "tail", after Graff. 

Rear of Polychoerus caudatus with 5 and 3 "tails", after 

2. Aphano stoma diversicolor , after Graff. 

3. Childia qroenlandica , from life. Actual size under one mm long. 

4. Anaperus qardineri , from life. 

5. Microstomum davenporti , from life. 

6. Monocelis sp. , from life. 

7. Plagiostomum sp., from life. 

8. Monoophorum sp., from life. 

9. Woodsholia lilliei (?)', from life. 


10. Gyratrix sp., with proboscis shown extended at right, from life. 

11. Macrostomum sp. , from life. 

12. Unidentified dalyellioid, from life. 

13. Bdelloura sp., probably Candida , from life. 

14. Syncoelidium pellucidum , from life, showing the diagnostic fusion of 
the hinder branches of the triclad gut. 

15. Procerodes wheatlandi , reconstructed outline from figures in Verrill. 

Abbreviations : 

a - adhesive papillae 

br - brain 

e - eyes 

eg - egg 

ge - genital pore 

m - mouth 

mp - muscles of proboscis 

o - ovary 

p - penis 

ph - pharynx 

pr - proboscis 

ps - penis stylet 

pu - purple cells 

s- sensory hairs 

sb - seminal bursa 

sp - sperm 

st - statocyst " 

su - sucker 

v - seminal vesicle 

vd - vas deference 

y - yolk gland 

Plate 4 




36 Turbellaria 

sensory hairs set in a thin area of epidermis and not on papillae. The worms 
from the Eel Pond should be good class material: they are large (up to 4 mm 
long) , can be reared in the laboratory, and regenerate readily. When kept in 
Syracuse dishes and fed small bits of annelid worms, they laid eggs which dev- 
eloped in about one week into small worms, like the adults except for undevel- 
oped reproductive structures. 

Order Tricladida 

Although marine triclads are in general uncommon, Woods Hole has several spe- 
cies. Bdelloura spp. (fig. 13) and Syncoelidium (fig. 14) are very easily found 
as commensals on the gill-books and around the leg bases of Limulus . The follow- 
ing annotated list may aid in the identification of the more common local triclads 
(figure references are to Plate 4) : 

Bdelloura Candida (Girard, 1850) . Abundant on Limulus ; the name is usually applied 
to any Bdelloura collected, although another species is also present. B. Can- 
dida reaches a length of 15 mm, and has 60-100 testicular sacs on each side of 

Bdelloura propingua Wheeler, 1894. This is described by William Morton Wheeler in 
his paper on Syncoelidium. B. propingua reaches a length of 8 mm, and has a- 
bout 170 testicular sacs on each side. 

Syncoelidium pellucidum Wheeler, 1894. Also found on Limulus but much less numer- 
ous than Bdelloura spp. Length 3 mm with about 15 testicular sacs on each side. 
The distinguishing feature of Syncoelidium is the fusion of the two posterior 
branches of the gut, and the lack of a distinct posterior sucker. 

Procerodes wheatlandi (Girard, 1850) is considered by Hyman (1944) "to be at best 
a geographic variant of P. littoralis " (Strom, 1768) which extends from New- 
foundland to Scandinavia. P. littoralis in turn may or may not be conspecific 
with Procerodes ( Gunda ) ulvae, which tolerates low and variable salinities and 
is well known as an experimental animal in European studies of osmoregulation. 
It is reported to be a small (5 mm long) dark animal with a pair of antero- 
lateral tentacles and 2 eyes (fig. 15). 

Order Polycladida 

Several species of polyclads are common at Woods Hole, and others occur less 
frequently. The key that follows is restricted to the commoner forms plus a couple 
that were easily included; consult the annotated list for forms that do not "key 
out" readily. Polyclads are in some cases very difficult to identify alive, and 
well cleared whole mounts or even sagittal sections are needed for serious work. 
In the key, only superficially visible chacteristics are employed; while this fac- 
ilitates an approximation to an identification, anyone doing experimental work on 
polyclads should be sure to fix material for determination by a qualified syste- 
matist before publication. 

For fixation, Hyman recommends hot seawater saturated with mercuric chloride, 
followed by thorough washing, dehydration in alcohol, and clearing in oil of win- 
tergreen. (For details consult Bull. Amer. Mus. Nat. Hist., 100 : 269 e_t seq . ) 

(Figure references on polyclads are to Plate 5) 

1. Tentacles absent 7 

1. Tentacles present (either "marginal" at front edge, or 

arising dorsally near the brain) 2 

Polycladida 37 

2. Tentacles formed by upfolded anterior margin; thin undulated 
margin; tentacles brownish on pale flesh colored ground; body 
10-12 mm long (fig .1) Prostheceraeus maculosus 

2. Tentacles dorsal, in region of brain (not always obvious 

when one looks directly down on living animal (figs. 2-4) 3 

3. Marginal eyes present on edge of body in addition to more 

centrally placed eyes 4 

3. Marginal eyes absent (but others may be present) 5 

4. Body somewhat elongated, with distinct pattern of white or 

yellowish and brown cross-bars Stylochus zebra 

4. Body broadly elliptical, of various shades of cream, yellow, 

gray, or brown (figs. 3,4) Stylochus ellipticus 

5. Body oval, tentacles do not contain scattered eyes 6 

5. Body elongated, widened anteriorly, pointed posteriorly; 
6-8 mm long; tentacles contain scattered eyes (fig. 2) ; 

body yellowish to pellucid with brownish spots Gnesioceros f loridana 

6. Generally found in mantle cavity of Busycon; color white 

Hoploplana inquilina 

6. Occasionally drifts in on Sargassum weed; color pattern of 

white reticulation on a brown ground Hoploplana qrubei 

7. Body yellowish to gray-brown; 8-12 mm long; slender, res- 
embling a fresh-water planarian; eyes near brain, typically 
4-5 on each side in a lengthwise row, and 2 set closely 

together a little to rear of each row (fig. 5) Euplana gracilis 

7. Eyes in 4 conspicuous clusters in area of brain (fig. 6); 

to be expected only north of the Cape Notoplana atomata 


Acerotisa baiae Hyman, 1940. Taken (Goodchild) at Lagoon Pond bridge (Hyman, 1952). 
The genus Acerotisa lacks tentacles and has 2 eye clusters. A. baiae is small 
(3 mm), translucent whitish. Not in key. 

Acerotisa notulata (Bosc, 1801) . A minute species that might occasionally drift 
into Woods Hole on Sargassum (Hyman, 1952). Not in key. 

Coronadena mutabilis (Verrill, 1873) . A rare southern form; status at Woods Hole 
uncertain. C. mutabilis lacks tentacles and has marginal eyes around anterior 
half of body; gray to yellowish brown; 18 by 5 mm. Not in key. 

Euplana gracilis (Girard, 1850) . Reported abundant in Eel Pond and among masses of 
sponges and hydroids on pilings. 

Eurylepta maculosa , see Prostheceraeus maculosus . 

G nesioceros f loridana (Pearse, 1938). A southern species, but recorded from eel 
grass in Woods Hole and Quissett Harbors, also on sandy bottoms at 8-10 meters 
(Hyman, 1939, 1940). There has been confusion about the name; it was once 
wrongly referred to Imogine oculif era , and in Hyman ' s good description (1939) 
it is called Gnesioceros verrilli . 

Gnesioceros sargassicola (Mertens, 1833). Has been taken from Sargassum in Vine- 
yard Sound (Hyman, 1939). Color pellucid with brownish spots; shape charact- 
eristic of genus, widened anteriorly, tapering to a pointed posterior end. 

Gnesiceros verrilli , see G. f loridana . 

Hoploplana qrubei (Graff, 1892) . Has been collected (Hadley) from Sargassum in 
Vineyard Sound (Hyman, 1939). 

Hoploplana inquilina (Wheeler, 1894) . Described by William Morton Wheeler as Plano - 
cer a inquilina , under which name its embryology was described by Surface in a 
well known study. Occurs locally in the mantle cavity of Busycon canaliculatum 

38 Polycladida 

but further south has been reported from Thais and Urosalpinx (Hyman, 1944) . 

Imoqine oculif era . see Gnesioceros f loridana . 

" Leptoplana " . used by E. B. Wilson (1894) in an often cited cell lineage study, is 
not a Woods Hole flatworm, but an unidentified species from Puget Sound, whose 
actual genus cannot be determined. 

Leptoplana variabilis . see Notoplana atomata . 

Notoplana atomata (0. F. Muller, 1776). Hyman (1939) calls this the commonest poly- 
clad of the North Atlantic shores, from northern Massachusetts to Scandinavia, 
but there is doubt that it extends south into inshore waters of Woods Hole. It 
has been reported from Nantucket. 

Planocera elliptica . see Stylochus ellipticus . 

Planocera inquilina . see Hoploplana ellipticus . 

Planocera nebulosa ; considered by Hyman (1944) a synonym of Stylochus ellipticus . 
P. nebulosa was used in the Invertebrate Zoology Course Keys during the 1940 ' s 
to refer to a greenish polyclad without marginal eyes, apparently following the 
usage in Pratt's Manual (p. 188). Verrill (1892, p. 472) points out thaf'P. 
nebulosa " has eyes that are difficult to see in the deeply colored living ani- 
mals. It is probably a color variant of S. ellipticus . 

Prosthecereaus maculosus (Verrill, 1892) . Reported "in some abundance" on pilings 
of Lagoon Pond bridge (Hyman, 1952) . This has been known as Eurylepta maculosa 
up until 1952, when an anatomical restudy by Hyman necessitated its transfer to 
Prosthecereaus . 

Stylochus ellipticus (Girard, 1850) . Under stones in shallow water and in tide 
pools. Feeds on oysters and barnacles (Hyman, 1940). 

Stylochus zebra (Verrill, 1882) . Easily recognized by color pattern. Usually col- 
lected from shells of Busycon occupied by the large hermit crab Pagurus polli - 
carus . but also found free living on rocks and pilings. 


Bresslau, E., 1933. Turbellaria, in_ Kukenthal and Krumbach (eds.) Handbuch der Zo- 
ologie , vol. II, Pt. l.(Wery complete general account with bibliography and 
most complete outline available of classification, including lists of genera) . 

Costello, D. P., et al. , 1957. Methods of Obtaining and Handling Marine Eggs and 
Embryos , MBL, Woods Hole (for Polychoerus and Hoploplana embryology and refer- 
ences) . 

Graff, L. von. 1882. Monographie der Turbellarian I. Rhabdocoelida - 2 vols. Leip- 
zig. (Many changes have made this classification out of date but the second 
volume of plates is worth looking over) . 

Graff, L. von, 1911. Acoela, Rhabdocoela and Allocoela des Ostens der Vereinigten 
Staaten von Amerika. Arbeiten aus den Zoologischen Institut zu Graz . IX Band, 
no. 8_: 321-428, 6 pi. (Of interest because of descriptions and drawings of Ana - 
perus, Childia and Woodsholia) . Same title, 1911, Z. wiss. Zool., 9J3: 1-108. 

Hyman, L. H. , 1939. Some polyclads of the New England coast, especially of the 
Woods Hole region. Biol. Bull. 76_: 127-52. 

, 1940. The polyclad flatworms of the Atlantic coast of the United 

States and Canada. Proc. U. S. Nat. Mus . 89_: 449-95. 

. 1944. Marine Turbellaria from the Atlantic coast of North America. 

Amer . Mus. Novitates, no. 1266: 1-15. 
, 1951. The Invertebrates, Vol. II. Platyhelminthes and Rhynchocoela. 

McGraw Hill, N. Y. (The most useful general reference on Turbellaria) . 
, 1952. Further notes on the turbellarian fauna of the Atlantic coast 

of the United States. Biol. Bull. 103_: 195-200. 

, 1959. Some turbellarians from the coast of California. Amer. Mus. 

Novitates, no. 1943: 1-17 (Synonymy of Childia). 
Meixner, J., 1938. Turbellaria, in Die Tierwelt der Nord- und Ostsee . Leipzig 

part IV: 1-146. (Mostly on biology and ecology of the turbellaria but includes 

Turbellaria 39 

an outline of classification which differs from that of Bresslau and has some 

advantages as a system) . 
Verrill, A. E. , 1892. Marine planarians of New England. Trans. Conn. Acad. Arts 

& Sci., 8: 459-520, Pi. 40-44. 
Wheeler, W. M. , 1894. Syncoelidium pellucidum , a new marine triclad. J. Morph. , 

9: 167-94, PI. VIII. (Contains also the description of Bdelloura propinqua ) . 

40 Chapter VII 


Nemertean worms are almost always easily recognized as such by their soft, elon- 
gated, narrow, highly contractile, unsegmented bodies, lacking setae and covered by 
cilia. A few species are common in the Woods Hole region, but about 30 may occur. 
The group is excellently discussed in Coe ' s "Biology of the Nemerteans of the Atlan- 
tic Coast of North America" (1943), which anyone making a serious study of the group 
must have at hand. However, identification by the beginner attempting to use the 
extensive keys in Coe ' s work is made difficult by the fact that the division of ne- 
merteans into orders is based in part upon such internal features as the arrange- 
ment of muscle layers in the body wall. The general descriptive features below may 
be useful in deciding the probable order in which to place an unknown specimen. 
Figure references on nemerteans are to Plate 5. 

Class ANOPLA: Mouth posterior to brain; proboscis not armed with stylets. 

Order Paleonemertea. In general, paleonemerteans are slender, soft, and ex- 
tensile, heads somewhat blunt, bodies not much flattened; ocelli and long- 
itudinal cephalic slits are lacking. 

Order Heteronemertea. Heads characteristically rather snakelike, with marked 
lateral slits (figs. 7, 12), but Parapolia and Zvqeupolia are exceptions. 
A small caudal cirrus (fig. 12) is found in Cerebratulus , Micrura , and 
Zvqeupolia . 

Class ENOPLA: Mouth anterior to brain; proboscis armed with one or more stylets 
(figs. 8-10) except in Bdellonemertea. 

Order Hoplonemertea. The stylets are diagnostic (flatten animals cautiously 
beneath cover slip or between microscope slides and examine by transmit- 
ted light) . 

Order Bdellonemertea. One species, commensal in mantle cavities of bivalve 

molluscs. Does not much resemble a nemertean, but has a leech-like form 
with posterior sucker (fig. 11). 


1. Free-living 3 

1. In mantle cavity of bivalves or in egg mass or gills of crabs 2 

2. In mantle cavity of bivalves; intestine convoluted and with- 
out diverticula; stylet apparatus absent; mouth and proboscis 
opening united; posterior sucking disk present (fig. 11); 

uncommon: Order BDELLONEMERTEA Malacobdella qrossa 

2. Among eggs or gills of crabs; proboscis rudimentary, with no 

accessory stylets Carcinonemertes carcinophila 

3. In marine or brackish waters 4 

3. In fresh water; with 4 or 6 ocelli; color reddish or pink- 
ish; length up to 20 mm Prostoma rubrum 

4. Generally small worms, with blunt, flattened heads; usually 
oblique cephalic grooves mark rear corners of head; ocelli 
usually obvious (4 or numerous) ; presence of proboscis sty- 
lets diagnostic Order HOPLONEMERTEA 5 

(Note: Prostoma and Carcinonemertes , keyed out above, are also 

in this order) 
4. Not referable to Hoplonemertea 15 

Nemertea 41 

5. Ocelli 4, set as corners of a square; small worm, often 

colorful 6 

5. Ocelli otherwise 11 

6. Body of slender cylindrical form; of firmer consistency 

than other nemerteans of similar small size (10-20 mm); color 
varies; head not demarcated from body; 4 ocelli in a square 
Oerstedia dorsalis 

6. Body of short, flattened form; head demarcated by incon- 
spicuous transverse grooves; 4 large, occasionally frag- 
mented, ocelli; worms small and variously colored Tetrastemma 7 

7. Body usually with more or less conspicuous longitudinal 

stripes 8 

7. Body lacking well defined longitudinal stripes 9 

8. Body rather slender, yellow, with 2 broad longitudinal 

brown stripes Tetrastemma elegans 

8. Body short and broad; usually green, with one or 2 longi- 
tudinal yellow stripes and 6 green stripes near tip of head 

Tetrastemma vittatum 

9. White or translucent with superficial flecks of white . . Tetrastemma wilsoni 

9. Yellow, rosy, red, or green 10 

10. Pale green or yellowish, head white or cream colored . . . Tetrastemma candidum 

10. Yellow or rosy, often spotted with brown; with band of 
dark pigment connecting the 2 ocelli on the same side of 

the head Tetrastemma vermiculus 

11. Ocelli extend posteriorly along lateral nerve cords beyond 
brain; basis of central stylet cylindrical and sharply trun- 
cated or concave at posterior end (fig. 9) Zygonemertes virescens 

11. Ocelli do not extend posteriorly beyond brain; basis of 
central stylet truncate-conical or pear shaped and usually 

rounded at posterior end 12 

12. Only one pair of ocelli, situated near tip of head; color 

of body orange-red Amphiporus bioculatus 

12. Ocelli 6 to 12 on each side of head 13 

13. Ocelli in a single row along each side of head (fig. 10); 
blood vessels bright red, and conspicuous in life; color 

of body usually pale yellow or rosy /Amphiporus cruentatus 

13. Ocelli in an irregular double row on each side or scat- 
tered 14 

14. Epidermis very thick and soft, secreting much viscid mucus 

when stimulated; movements of body often leech-like .... Amphiporus griseus 

14. Epidermis thin and firm, secreting but little mucus when 

stimulated; moves by creeping Amphiporus ochraceus 

Note: The following dichotomy involves separating the orders PALEONEMERTEA 
and HETERONEMERTEA . The mouth in both is posterior to brain; proboscis 
stylets are absent. 

15. Not possessing ocelli, or longitudinal cephalic grooves, or 
caudal cirrus; worms slender and not over 150 mm long 
probably Order PALEONEMERTEA 16 

15. Any large local nemerteans; any nemerteans with deep longi- 
tudinal cephalic slits, or with a caudal cirrus (but this is 
often broken off), o_r with ocelli but without stylets 
probably Order HETERONEMERTEA 18 

42 Nemertea 

16. Bodies filiform (lengths up to 150 mm, diameter up to one mm 17 

16. Body much flattened in posterior region (width 2-5 mm) ; 
head changeable in shape, broader than adjacent body, flat, 
rounded or emarginate anteriorly; color pale reddish or yel- 
lowish Car i noma tremephoros 

17. Very slender, filiform body; head broad; mature worms up 
to 25 mm long; color whitish; forms delicate mucoid tubes 

Tubulanus pellucidus 

17. Very slender, filiform body; head long, pointed; mouth far 
back of brain; length up to 100 mm; characteristically coils 

body into a close spiral Procephalothrix spiralis 

fclote: A worm of similar appearance, but not tending to con- 
tract into a spiral, is Cephalothrix linearis , not reported 
south of Cape Cod) . 

18. Without longitudinal cephalic grooves 19 

18. With longitudinal cephalic grooves 20 

19. Caudal cirrus present; head narrow, without oblique ceph- 
alic grooves Zygeupolia rubens 

19. Caudal cirrus absent; head broad; with oblique cephalic 

grooves Parapolia aurantiaca 

20. Caudal cirrus absent; body long and slender, filiform in 
some species, rounded or flattened in others, very contrac- 
tile; ocelli present in most species Lineus 21 

20. Caudal cirrus present; body not very slender, ocelli pre- 
sent or absent 23 

21. With conspicuous median dorsal stripe, but without trans- 
verse markings; reddish brown or olive, with median dorsal 
stripe of white or yellow extending whole length of body 

and head (fig. 7) Lineus bicolor 

21. Without conspicuous median dorsal stripe 22 

22. Head rather broad, cephalic grooves short; body contracts 

by shortening and thickening -- not by coiling in spiral .... Lineus ruber 

22. Head narrow, cephalic grooves long; body contracts by 

coiling in spiral Lineus socialis 

23. Body firm, long and ribbon-like, sometimes very large; much 
flattened in intestinal region, with thin lateral margins 
and well adapted for swimming; body less contractile than in 
other genera; mouth large and elongated, ocelli absent (fig. 

12 ) Cerebratulus lacteus 

23. Body slender, flattened in intestinal region, but with lateral 
margins not thin; incapable of swimming; mouth small and round; 

ocelli present or absent Micrura 24 

24. Head with a row of 4-6 ocelli on each side Micrura af finis 

24. Head without ocelli 25 

25. Color of body red or reddish 26 

25. Color of body whitish or pale yellowish; may show a tinge of 

red or orange anteriorly Micrura albida 

26. Deep red or purplish red; common Micrura leidy i 

26. Pale red, yellowish-red, or brownish-red Mi crura caeca 

Nemertea 43 



Order Paleonemertea 

C arinoma tremaphoros Thompson, 1900. Common in sand, clay, mud, or under stones. 
Cepha lothr ix linearis (Rathke, 1799) . Not reported south of Cape Cod. 
Proce p halothrix spir alis (Coe, 1930) . Common. 
Tub ulanus pel lucidus (Coe, 1895) . Not very common. 

Order Heteronemertea 

Cerebratulus lacteus (Leidy, 1851) . The commonest large nemertean; classical embry- 
ological material. 

Lineus bicolor Verrill, 1892. Common; usually subtidal. 

Lineus ruber (O. F. Muller, 1771). Often in low or variable salinity; color varies. 

Lineus socialis (Leidy, 1855). Common; often gregarious. 

Micrura aff inis (Girard, 1853). A northern species; below 10 m off Martha's Vine- 

Micrura albida Verrill, 1879. Not reported south of Cape Cod. 

Micrura caeca Verrill, 1895. Under stones or in sand. 

Micrura leidyi (Verrill, 1892) . Common in protected bays. 

Parapolia aurantiaca Coe, 1895. 

Zygeupolia ru bens (Coe, 1895). Abundant in sand in bays and estuaries. 

Order Hoplonemertea 

Amphiporus bioculatus Mcintosh, 1873. Common, subtidally in Vineyard Sound. 

Amphiporus cruentatus Verrill, 1879. Common locally in Woods Hole area. 

Amphiporus griseus (Stimpson, 1857). Occasional at Woods Hole. 

Amphiporus ochraceus (Verrill, 1873) . Common in Woods Hole area. 

Carinonemertes carcinophila (Kolliker, 1845). 

Oerstedia dorsa lis (Abildgaard, 1806) . Locally abundant, among growth on rocks and 

pilings . 
Ototyph loneme rtes pellucid a Coe, 1943. A minute form. Not in key . The genus is 

unique in possessing statocysts. 
Prostoma rubrum (Leidy, 1850). In fresh water swamps and ponds. Coe reports a 

green variety in a cedar swamp near Woods Hole. 
T etrastemma candid um (Muller, 1874). Common. 
T etrastemma eleqans (Girard, 1852) . Occasional. 
Tetrastemm a vermiculus (Quatref ages , 1846). Common. 

T etrastemma vittatum Verrill, 1874. Occasional, in protected muddy situations. 
Tetrastemma wilsoni Coe, 1943. Among Bryozoa, sponges, etc. on pilings. 
Zygonemertes virescens (Verrill, 1879) . Common. 

Order Bdellonemertea 

Mala cobdella g rossa (0. F. Muller, 1776). In mantle cavity of Mya . Mercenaria , 
O stre a and other bivalves. Not common around Woods Hole. M. obesa and M. 
mercenaria of Verrill are synonyms. 


Coe, W. R., 1943. Biology of the Nemerteans of the Atlantic Coast of North Amer- 
ica. Trans. Conn. Acad. Arts & Sci., 3_5: 129-328. 

Verrill, A. E. , 1892. The marine nemerteans of New England and adjacent waters. 
Trans. Conn. Acad. Arts & Sci., 8: 328-456, pi. 33-39. 

McCaul. W. E. , 1963. Rhynchocoela : Nemerteans from marine and estuarine waters of 
Virginia. J. Elisha Mitchell Sci. Soc . , 79: 111-124. 


Plate 5 


Polycladida (figs. 1-6), Nemertea (figs. 7-12), Sipuncul- 
oidea (figs. 13, 14), Nematomorpha (figs. 15-18). Figs. 
11, 12, 15 by Mrs. Emily Reid; figs. 13, 14, 16-18 by 
Bruce Shearer. Scales various. 

Fig. 1. Prostheceraeu s maculosus . outline to show marginal tentacles; 
note cerebral and tentacular eyes. 

2. Gnesioceros f loridana , simplified after Hyman (1939) to show body 
outline, cerebral and tentacular eyes. 

3. Stv l ochus ellipticus , simplified, showing tentacles and eyes. 

4. Stvlochus ellipticus . viewed from side to show dorsal tentacles, and 
marginal, tentacular, and cerebral eyes. 

5. Euplana gracilis , from life, showing pattern of cerebral and 
"tentacular" eyes. Note that actual tentacles are absent. 

6. Notoplana atomata, outline of body and pattern of eyes, simplified 
after Hyman (1939). 

7. Lineus bicolor (Heteronemertea) , head in dorsal view, from life, 
showing left cephalic slit and eyes. 

8. Amphiporus ochraceus (Hoplonemertea) , proboscis stylets as seen in 

a worm flattened on a slide; central stylet on pear shaped basis and 
accessory stylets in 2 lateral pouches. Drawn from a photo taken by 
Dr. W. E. McCaul. 

9. Zvqonemertes virescens , head of older animal with many eyes, stylet 
with truncated basis. After Coe. 

10. Amphiporus cruentatus , head and stylet. After Coe. 

11. Malacobdella grossa . After Verrill. 

12. Cerebratulus lacteus (Heteronemertea), whole animal with head in 
ventral view, showing mouth, left cephalic slit, proboscis pore an- 
teriorly, and caudal cirrus. After Verrill. 

13. Golfingia gouldi , from life; one with extended introvert showing 
tentacles, other with introvert withdrawn; about half natural size. 

14. P hascolion strombi , with extended introvert; much enlarged (scale 
bar equals one mm) . 

15. N ectonema agilis , impression of the worm as seen swimming. About 
natural size. 

16. Nectonema agilis , anterior end of living specimen much contracted 
after shedding eggs. Note 2 rows of natatory bristles. 

17. Nectonem a agilis , anterior end of relaxed, living specimen, prior 
to shedding of eggs. 

18. Nect onema agilis , posterior end of the above specimen. 

Plate 5 

46 Chapter VIII 



Here are included a number of classes (sometimes regarded as phyla) of pseudo- 
coelomate types, most of which are either absent from the local marine fauna (Pria- 
pulida) or of such small size, diversity, and great number of species as to make 
their inclusion in these keys impractical (Rotifera, Gastrotricha, Kinorhyncha, 
Nematoda) . The best general reference on aschelminth groups, and the first source 
of information when one encounters an unfamiliar member of them, is Hyman's "The 
Invertebrates", Vol. Ill: "Acanthocephala, Aschelminthes, and Entoprocta" , 1951. 

The remaining class, Nematomorpha or Gordiacea, has one marine order, Nectone- 
matoidea, represented by the single genus Nectonema . N. agilis Verrill, 1879 can 
be taken at a night light near the breakwater at the U. S. Fish and Wildlife lab- 
oratory in Woods Hole. It resembles a whitish "horsehair snake" provided with a 
double row of fine natatory bristles (Plate 5, figs. 15-18) with a body up to one 
mm in diameter and 20 cm long, and swims with a remarkable rapid, undulatory mo- 
tion. The young stages are reported to be parasitic in several species of true 
crabs and hermit crabs on the coast of France, but the life cycle at Woods Hole 
has never been observed. Hyman (loc. cit) provides the most helpful general ac- 
count of Nectonema . 


The unsegmented wormlike creatures formerly lumped as "Gephyreans" include 
three groups now commonly accorded phylum status: Priapulida, Sipunculoidea, and 
Echiuroidea (the last, however, is again joined to the annelids by Hyman). At 
Woods Hole, only the Sipunculoidea are represented, intertidally and in shallow 
water, by Golf ingia g ouldi (Pourtal^s, 1851) (Plate 5, fig. 13), and by Phasco - 
lion strombi (Montagu, 1804) (fig. 14), a very small worm from 10 meters and 
deeper off the Elizabeth Islands near Woods Hole, living in deserted gastropod 
shells or in tubes of Hydroide s , Pectinari a , Hyalinoecia , or other worms. 

Golf ingia oouldi has long been known as Phascolosoma gouldi and since there 
is no question about the species, experimental biologists should be careful to 
use the specific name to avoid confusion with certain common sipunculids to which 
the generic name Phascolosoma has now been transferred (e.g. Phascolosoma agassizi , 
formerly Physcosoma, of the West Coast, a species far less euryhaline than the 
Woods Hole Golf ingia gouldi ) . The reasons for the unfortunate shift of names 
are, in brief: Phascolos oma was established as a genus in 1828 by Leuckhart, 
with type species Phascolosoma granulatum of the Mediterranean. Later, in 1866, 
Quatref ages broke Leuckhart ' s genus into subgenera and used P_. granulatum as the 
type of a new subgenus Phymosomum . This was amended to Phymosoma by Salenka, 
and raised to generic rank in 1883. Phymosoma , being found preoccupied, was 
changed in 1897 to Physcoso ma. This whole procedure was in clear violation of 
nomenclatorial rules, according to which the type species of Phascolosoma should 
have retained its name. In 1952, Fisher put P hyscosoma back into its proper 
category of Pha scolosoma , but this left the group long known as Phascolosoma with- 
out a name. The "next available" name for Phascolosoma gouldi was Golf ingia , and 
this solution seems to have been generally accepted. 


Fisher, W. K. , 1952. The Sipunculid Worms of California ... , Proc. U. S. Nat. 

Mus. , 102 : 371-450. 
Gerould, J. M. , 1913. The sipunculids of the eastern coast of North America. 

Proc. U. S. Nat. Mus., 4±: 373-437. 

Chapter IX 47 


The Annelida or segmented worms have, as a phylum, shown remarkable evolutionary 
success in adapting to the marine, fresh water, and terrestrial habitats. Among the 
classes of Annelida, the Polychaeta are the typical marine representatives, although 
members of the other classes are to be found as well. 

Class Hirudinea (Leeches) 

Leeches are far less common in the sea than in fresh water, except for one fam- 
ily, Ichthyobdellidae. In the Woods Hole region the following may be encountered 
attached to fishes; the group, however, needs restudy in this area: 

Branchellion raveneli (Girard, 1851) . On the sting ray, Myliobatis fremmviller . 
Easily recognized by the leaf like gills along the sides of its body. 

Ichthvobdella funduli Verrill, 1872. Reported on Fundulus heteroclitus . Body 
smooth, annulated, greenish; 2 large and 2 small eyes. 

Ichthyobdella rapax (Verrill, 1873). On the summer flounder, Paralichthys dentatus . 
Body in extension long, slender, rounded, 3-4 cm, long, dark olive with row of 
rectangular white spots along each side. 

Trachelobdella vividus (Verrill, 1872). Body with two distinct regions; narrow an- 
teriorly and wide posteriorly; brown to purplish; about 2.5 cm long. On skates 
and rays; apparently not common. 

Reference: Knight -Jones, E. W. , 1962. The systematics of marine leeches. App. B. 

in "Leeches (Hirudinea) Their Structure, Physiology, etc.", K. H. Mann, Pergamon 

Class Oligochaeta 

Oligochaete annelids are characteristically fresh water or terrestrial; however 
there are some 10 species reported from marine or brackish waters of the New England 
coast, and some are very commonly encountered in shore collecting in the region of 
Woods Hole. Identification requires study of the internal anatomy, and no key is at 
present available other than that given by Moore: 

Moore, J. P., 1905. Some marine Oligochaeta of New England. Proc. Acad. Nat. Sci. 
Phi la., 5J7: 373-399, pi. 32-33. 

Class Archiannelida 

Archiannelids have, as the name implies, been regarded as possibly primitive 
forms, although many zoologists now prefer to regard them as secondarily simplified 
and very small derivatives of various polychaete families. They are characteristic- 
ally tiny marine worms, in sand, mud, or among algae, with few setae, parapodia re- 
duced or lacking, and often with external ciliation persisting in the adult. Be- 
cause of their very small size, archiannelids may be more common than is generally 
supposed; however, only one species of Dinophilus is collected regularly at Woods 
Hole. The morphology of Dinophilus (although not of a species reported from Woods 
Hole) is described by Nelson (1907). 

The species of archiannelids reported from the Woods Hole region include: 

Dinophilus gardineri A. Moore, 1899. Reddish in color; 6 segments; body with cil- 
iated rings but with no setae; occurs in brackish water such as Lillie's Ditch 
and the Eel Pond at Woods Hole. Figure 49 (Plate 8) is probably of this species. 

Dinophilus pygmaeus Verrill, 1892. Described as whitish, of 5 body segments, about 
0.7 mm long, and occurring on pilings at Woods Hole. 

Chaetogordius canaliculars J. P. Moore, 1904. Slender, filiform, 30 mm long, with 
a pair of small tentacles; posterior 10-12 segments with setae. 

48 Annelida 


Nelson, A. J., 1907. The morphology of Dinophilus conklini n. sp. Proc. Acad. Nat. 
Sci. Phi la., 59: 82-143, PI. 12-13. 

Class Polychaeta 

Polychaetes of the Woods Hole area are extremely numerous and diverse. Their 
treatment in a key is rendered difficult by the fact that, although the polychaetes 
can be divided into a large number of families, there is no real agreement as to how 
these families may be grouped into orders. The concept of a Family in the Polychaeta 
is important and practical for the student to grasp, for the reason that the basic 
structural features that determine a family also determine a way of life. With a 
little practice, most families may be readily recognized on sight, and many problems 
of keying are thus reduced to the level of genera and species, where even the spec- 
ialist may have to make a close examination. The problem of identifying certain fam- 
ilies is simplified by attention to the very characteristic tubes formed by some. 

There is still no completed monograph of the polychaetes of the Woods Hole region, 
but this need will be met by the publication of Pettibone ' s 'Warine Polychaete Worms 
of the New England Region", of which Part 2 is now in preparation. Until this appears, 
the most useful complete work is the two volumes by Fauvel; these deal with the poly- 
chaetes of France, but are exceedingly useful in identifying families and genera, as 
well as some of our local species. There also exist Dr. Pettibones mimeographed pre- 
liminary check list and key to Polychaeta of New England; copies of this are available 
in the invertebrate class laboratory, and should be consulted for forms which do not 
clearly "fit" in the simpler key presented below for general use. This key is not in- 
tended to satisfy the specialist, but rather to bridge the gap between the general 
biologist and the polychaete systematist. It will probably satisfy neither group, and 
as a result of this irritation, it is hoped that improved versions will arise. 

Much remains to be learned. Polychaete families are of convenient dimensions for 
individual student projects, and offer much of interest to the comparative physiologist 
and ecologist. The following key is frankly a preliminary attempt, and will need much 
revision in the future. We are grateful to Dr. Marian Pettibone for much help and ad- 
vice in the preparation of the key and check list. 

A Note on Polychaete Terminology 

There is no generally accepted scheme for dividing the Polychaeta into orders. 
One method has been to recognize one order of more or less sedentary forms (Poly- 
chaeta Sedentaria) and another of free living types (Polychaeta Errantia) . Actually, 
such a division is arbitrary, since the families of polychaetes present a spectrum of 
adaptions, ranging from free swimming types, through crawlers, burrowers and temporary 
tube builders, to those which construct fixed tubes which they never leave. In prac- 
tice it is more useful to recognize the principal family types, such as nereid, poly- 
noid, spionid, syllid, sabellid, serpulid, etc. 

A typical free living polychaete such as Nereis (which will serve as an example, 
although members of other families differ greatly in detail) has paired locomotor ap- 
pendages or parapodia (sing., parapodium) (Pi. 7 , figs. 12-15), each composed of an 
upper lobe, the notopodium, and a lower, neuropodium. Each lobe typically contains 
a bundle of slender chitinous setae (sing, seta) which project from the parapodium, 
together with a larger dark spine known as the aciculum ( pi . acicula) . The shape, 
size, number, and position of the setae are of importance in classification. Arising 
from the base of the notopodium above and the neuropodium below, there are often slen- 
der flexible outgrowths, the dorsal cirrus (pi. cirri) and ventral cirrus, respectively. 
The notopodium consists of a dorsal lobe or ligule and a middle lobe, between which are 
the aciculum and the setae; the neuropodium consists of a neuroacicular lobe provided 
with setae and an aciculum, and a ventral lobe. In many polychaetes, gills or branch- 

Polychaeta 49 

iae, made conspicuous by the red or green blood within, arise from or near the para- 
podia in certain parts of the body. 

Setae vary widely in form, and furnish very precise characters for the determina- 
tion of species. The use of such characters has been avoided where possible in the 
key but is necessary in some cases. Common types and their names should be recognized. 
Plate 6 gives some idea of the wide variety of types. First we may distinguish sim- 
ple setae (figs. 1-20, 29-42) from composite or jointed setae (figs. 21-28). Long 
slender simple setae are spoken of as capillary setae (fig. 6) . The tips of setae, 
whether simple or composite, may be entire, bifid (fig. 8), or trifid. If bent like 
a sickle they are termed falcate (fig. 11) ; if flattened like an oar blade, limbate 
(fig. 10). Some simple setae have stubby, bent, usually bifid ends. These are spo- 
ken of as hooks or crochets (figs. 29-32). These are usually relatively stout setae 
and grade into short, broadened types known as uncini (figs. 33-42), usually set in 
close rows, which are especially characteristic of the Sedentaria. 

Composite setae may be multiarticulate, as in the long bristles of some flabel- 
ligerids (fig. 21), but are characteristically two-jointed, composed of a shaft and 
a blade (figs. 22-28). This blade in turn may have various shapes and may itself be 
a crochet (fig. 28) . The blade rests in a notch in the end of the shaft. If the two 
sides of the notch are equal, it is spoken of as homogomph (fig. 27) ; if unequal, 
as heterogomph (fig. 26). Finally setae, either simple or compound, may be embedded 
at their tips in a clear matrix and are then spoken of as hooded. Thus, we may have 
simple hooded crochets (figs. 20, 29), or composite hooded crochets (fig. 28). 

The heads of polychaetes are exceedingly diverse. As an example of the head of 
an errant polychaete we may take the head of Nereis (Plate 7 , figs. 8-11). This 
consists of a preoral prostomium, provided at its anterior margin with a pair of small 
antennae, and at its sides with paired, fleshy, biarticulated palps (palpi) . The seg- 
ment just behind the prostomium is the peristomium (in Nereis it represents a fusion 
of two segments). It bears, in Nereis , four pairs of peristomial cirri (tentacular 
cirri) on short stalks at its anterior margin. The usage of the terms palp, antenna , 
and cirrus varies greatly. Antennae, unless otherwise specified, are usually dorsal 
or marginal on the prostomium (the terra tentacle may also be used). Palpi are usually 
associated with the mouth and tend to be lateral or ventral to the prostomium and bor- 
dering the anterior margin of the mouth. However, certain dorsal structures, especial- 
ly if these are large, elongated, grooved, or prehensile (as in the spionids) are fre- 
quently called palps. The term cirrus is usually applied to structures arising dor- 
sally or ventrally on the parapodia, whereas comparable structures on the anterior 
part of the body, if elongated, may be designated tentacular cirri (or peristomial ten- 
tacles) . Tentacle is a very general term and is used to signify any of a variety of 
elongated sensory or feeding structures, usually on the head. 

The prostomium of Nereis bears two pairs of eyes; other polychaetes may have one 
pair or none, and in some there are numerous eyespots scattered on the peristomium or 
even on the tentacles or sides of the body. 

The first parapodia of Nereis are borne on the segment behind the peristomium. 
The first two pairs of parapodia are uniramous, that is, each contains a single acicu- 
lum and setal fascicle (bundle). All succeeding parapodia are biramous . In other 
polychaetes the parapodia vary, some having only uniramous, others only biramous pa- 
rapodia, some with both types, some with parapodia greatly reduced. 

In most free living polychaetes the pharyngeal region may be everted (pushed for- 
ward and turned inside out) to form a proboscis, which often bears stout jaws and small 
horny teeth (paragnaths) . A study of the arrangement of paragnaths is a necessary step 
in the identification of nereids; in other families the pattern is less complex, but 
may be referred to. 

The sedentary polychaetes depart widely from the body form which characterizes 
free living types. Prostomium and eyes are often reduced, proboscis and jaws may be 
absent, and the anterior end, especially in types dwelling in fixed tubes, greatly 

50 Polychaeta 

elaborated for feeding and respiration. In sabellids and serpulids the peristomial 
cirri (or tentacles) form a "branchial crown" of feather-like "gills", which serves 
both for feeding and respiration. Cilia pass water between the branches of the plumes 
and transport food, entangled in mucus, down to the mouth. In other forms such as 
terebellids, the peristomial tentacles are long, filamentous, and extensile, serving 
to bring in food by ciliary action in a groove running along each filament; just be- 
hind the head there arise blood-filled branchiae which serve in respiration. Para- 
podia in tubicolous (tube dwelling) polychaetes tend to be small and are provided 
with rows of hooklike setae (uncini) for gripping the sides of the tube. Uncini 
may be set on a low mound or uncinigerous torus. Special glands may secrete tube- 
forming material. One or more peristomial cirri may, in serpulids, which form a 
rigid calcareous tube, be modified to a pluglike operculum that can block the tube 
entrance. The body in tube dwelling forms is often divisible into a more anterior 
and specialized thorax and a less specialized posterior abdomen. The thorax may 
bear anteriorly a collar, and this may be extended rearward to the posterior end of 
the thorax as a pair of folds, the thoracic membranes. The preceeding accounts are 
of extreme types, free living and tubiculous respectively, and can convey but a poor 
idea of the actual diversity of pattern that characterizes the numerous annelid 
families . 

The key that follows begins as a key to separate families, and a study of it will 
give an impression of the characteristics of each family type. Since the ability 
to recognize families in the field is of great practical value to the zoologist, the 
student should early learn the family types commonly encountered. Generic and spe- 
cific identification requires more careful study. (For setal types see Plate 6; other 
figure references are to Plates 7-10) . 


I. Instead of presenting only a pair of choices, this key presents several choices 
at certain steps. Figure references in Keys are to Plates 7-10. 

II. Effective use of this key requires practice. We suggest that the beginner, with 
the advice of the instructor, run an assortment of well known species through the 
key step by step, in order to acquaint himself with the terminology. The tendency 
of beginners to skip steps when the worm at hand is known interferes with learn- 
ing to use the key. 

Ill . In most families only those species are keyed out which are commonly collected. 
The student is referred to the more detailed key of Dr. Pettibone for the rarer 
species, or those in which identification must be based upon more details than 
given here. 

IV. The keys to members of families having more than one common or readily identifi- 
able species will be found under the separate families, together with annotated 
lists of species reported from this region. No attempt has been made to key out 
all the rare or deeper water species. 

V. Unidentifiable and otherwise "interesting" polychaetes may be sent to Dr. Marian 
Pettibone, Division of Marine Invertebrates, U. S. National Museum, Washington 
25, D. C. 

1. Dorsal surface with overlapping scales (elytra, fig. 1), flat- 
tened setae (paleae, fig. 22), or overlain by a thick felty layer 2 

1. Dorsal surface not with elytra, paleae, or felt 4 

(Key continued on page 60) 

Plate 6 

21 22 23 25 26 28 29 32 


37 42 

52 Plate 7 


Polynoidae, Euphrosinidae, Spintheridae, Ampharetidae, 
Flabelligeridae, Nereidae, Onuphidae, Eunicidae, Scali- 
bregmidae, Dorvilleidae , Spionidae. 

Note: In this and in Plates 8, 9, and ID, objects are not 
to any scale size. Figure sources are: (F) Fauvel, 
(B) Berkeley and Berkeley, (P) Pettibone, (S) from 
specimens . 

Fig. 1. Lepidonotus squamatus . Polynoidae, showing 12 pairs 
of elytra (singular is elytron) (S) . 

2. Head of Harmothoe imbricata (F) . 

3. Euphrosine borealis , dorsal view of anterior end (P) . 

4. Spinther citrinus , dorsal view of anterior end (P) . 

5. Hypaniola grayi , Ampharetidae (S) . 

6. Anterior end of Polydora sp. (Spionidae), showing modified 
fifth setiger (after Verrill) . 

7. Pherusa af finis , Flabelligeridae, anterior end in lateral view (S) 

8. Nereis ( Neanthes ) succinea , head and everted proboscis in 
dorsal view (S) . 

9. Platervnereis meqalops . proboscis of male in ventral view 

10. Nereis ( Neanthes ) arenaceodonta . proboscis, ventral view (S) . 

11. Head and proboscis of same, dorsal view (S) . 

12. Parapodium from posterior part of body of N. succinea (S) . 
In this and figs. 13-15, parapodia are from left side, 
viewed from front. Acicula are shown in black: omitted 

in figs. 13-15. 

13. Parapodium from mid-body of Nereis ( Nereis ) pelagica , show- 
ing 2 ligules in notopodium (S) . 

14. Same, from N. ( Neanthes ) virens . showing 3 ligules in noto- 
podium. (S) . 

15. Same, from N. (Neanthes ) arenaceodonta (S) . 

16. Head of generalized onuphid, showing pair of nuchal ten- 
tacles on peristome (P) . 

17. Eunice pennata , Eunicidae. diagram of head (P). 

18. Marphysa sanguinea , same (S) . 

19. Marphysa belli , same (P) . 

20. Scalibregma inf latum , Scalibregmidae, same (P) . 

21. Stauronereis rudolphi , Dorvilleidae (F) . 

Plate 7 

54 Plate 8 


Chrysopetalidae, Syllidae, Terebellidae, Cirratulidae 

Fig. 22. Dvsponetus pygmaeus (Chrysopetalidae), dorsal view of 
whole animal showing dorsal fans of setae (P) . 

23. Syllis gracilis , head (P) . 

24. Same, parapodium (P) . 

25. Same, compound setae. 

26. Same, heavy bifurcated simple setae (P) . 

27. Amblyosvllis formosa (European) to show nuchal epaulettes (F) 

28. Odontosvllis fulgurans , head showing nuchal hood (F) . 

29. Same, armature of proboscis (F) . 

30. Streptosyllis , diagram of parapodium with knobbed aciculum 

31. Eusyllis lamelligera, head (F) . 

32. Same, armature of proboscis (F) . 

33. Brania clavata , head (P) . 

34. Exogone hebes , head (P) . 

35. Exogone dispar , head (P) . 

36. Sphaerosyllis e rinaceus . head (P) . 

37. Autolvtus prismaticus . head (P) . 

38. Arborescent (branching) gill of a terebellid (S) . 

39. A terebellid gill of unbranched filaments (B) . 

40. Head of adult Nicolea venustula , simplified (P) . 

41. Head of Pista maculata , simplified (P) . 

42. Head of Pista cristata . simplified (B) . 

43. Head of Terebellides stroemi , simplified (B) . 

44. Gill-like branching parapodium of Enoplobranchus sanguinea 
(note setae) (P) . 

45. Dodecaceria coralii , Cirratulidae, whole animal (S) . 

46. Cirratulus cirratus , anterior end (P) . 

47. Tharyx , sp., anterior end (F) . 

48. Chaetozona setosa , head (P) . 

49. Dinophilus sp., Archiannelida, drawn from life by Dr. 
Louise Bush. 

Plate 8 

56 Plate 9 


Sabellidae, Phyllodocidae, Serpulidae, 
Maldanidae, Hesionidae, Paraonidae 

Fig. 50. Fabricia sabella , whole worm, dorsal view (F) . 

51. Chone infundibuliformis . simplified, anterior end, view of mid-dorsal slit 
in collar (F) . 

52. Sabella crassicornis , anterior end viewed from left; note 4-lobed collar (P) . 

53. Sabella microphthalma , collar region dorsal view; irregular rows of eyes (S) . 

54. Same, ventral view; note absence of lateral slits (S) . 

55. Potamilla renif ormis , dorsal view of collar region, showing mid-dorsal and 
lateral notches; "compound eyes" on branchial filaments (S) . 

56. Eteone sp., anterior end (P) . 57. Phyllodoce sp., anterior end (S) . 

58. Notophvllum americanum , anterior end (P) . Note nuchal lappets at rear of 
prostomium, and large imbricated dorsal cirri. 

59. Eulalia viridis . anterior end (P) . 

60. Eumida sanquinea , anterior end (P) . 61. Paranaitis speciosa , anterior 

end (P) . 

62. Spirorbis ( Dexiospira ) spirillum , dextrally coiled tube with end elevated (F) 

63. Same, dextral, flat coil (P) . 

64. Spirorbis ( Paradexiospira ) violaceus , dextral tube (F) . 

65. Spirorbis ( Laeospira ) borealis , sinistral tube (F) . 

66. Operculum of above (P) . 67. Spirorbis ( Laeospira ) granula - 

tus , sinistral tube (P) . 

68. Operculum of 67 (P) . 69. Operculum of Filograna implexa 


70. Operculum of Hydroides ( Eupomatus ) uncinata (P) . 

71. C lvmenella torquata , anterior end viewed from right, showing collarette on 

4th setiger (P) . 

72. Posterior end of same, showing anal funnel (P) . 

73. Posterior end of Maldanopsis elongata (P) . 

74. Posterior end of M aldane sarsi (P) . 

75. Nephtys incisa , head end (P) . 76. Nephtys bucera , head end (P) . 

77. Aglaophamus circinata , parapodium with branchia curved inwards (P) . 

78. Nephtys caeca , parapodium from middle region, posterior view, showing 
branchia curved outward (P) . 

79. Podarke obscura , head (S) . 80. A ricidea jeffreysii , head (B) . 

Plate 9 

58 Plate 10 


Spionidae, Goniadidae, Glyceridae, Arabellidae, 
Lumbrineridae, Orbiniidae, Oweniidae, Pectinariidae, 

Sabellariidae . 

Fig. 81. Polydora ligni , head with occipital antenna (P) . 

82. Spiophanes bombyx , prostomium (P) . 83. Scolecolepides viridis , head (P) . 

84. Dispio uncinata , head (P) . 85. Scolelepis squamata , head (P) . 

86. Spio filicornis , head (P) . 87. Hooded neuropodial crochet of Spio 

f ilicornis (P) . 

88. Goniada maculata , head with extended proboscis, showing chevrons (Mcintosh). 

89. Glycera capitata , anterior parapodium seen from rear, showing single post- 
setal lobe (P) . 

90. Glycera dibranchiata , parapodium seen from rear, showing 2 postsetal lobes 
and 2 branchiae (P) . 

91. Glycera americana , parapodium from middle body, seen from rear, showing the 
branchiae extended (P) . 

92. Ophioglycera gigantea (Goniadidae), anterior uniramous parapodium seen from 
front, showing presetal lobe (P) . 

93. Same, posterior biramous parapodium, seen from front, also with bilobed pre- 
setal lobe (P) . 

94. Lumbrineris fragilis , capillary 95. Same, hooded seta (P) . 
limbate setae (P). 

96. Ninoe nigripes , hooded seta (S) . 97. Same, branchia bearing para- 
podium seen from front (P) . 

98. Lumbrineris acuta , head (P) . 99. Lumbrineris tenuis , head (P) . 

100. L. tenuis , posterior parapodium (P) . 101. Arabella iricolor , head (P) . 

102. Driloneris longa , anterior para- 103. Same, posterior parapodium (P) . 
podium (P) . 

104. Orbinia ornata , right hand parapodium in thoracic region, seen from rear, 
showing palisaded neurosetae and postsetal row of papillae (P) . 

105. Scoloplos robustus , abdominal parapodium, showing undulate subpodial flange (P) 

106. Scoloplos fragilis , abdominal parapodium showing incised subpodial lobe (P) . 

107. Owenia fusiformis , anterior end seen from side, showing slashed branchial 
membrane (P) . 

108. Pectinaria gouldii , seen in dorsolateral aspect; note operculum of golden 
paleae (S) . 

109. Sand tube of Pectinaria , about natural size (S) . 

110. Sabellaria vulgaris , living specimen removed from tube (S) . 

Plate 10 

60 Polychaeta 

2. Dorsal surface with a felty layer over elytra; "sea mice" 


2. Dorsal surface with simple, exposed elytra; "scale worms" 3 

2. Dorsal surface more or less concealed by notosetae in fan- 
shaped groups or transverse rows; minute worms CHRYSOPETALIDAE 

The only species reported locally is Dysponetus pygmaeus 

Levinsen, 1879; dredged; up to 3 mm long (fig. 22). 

3. With filiform dorsal cirri on non-elytra bearing segments 


3. Without filiform dorsal cirri on non-elytra bearing segments 


The only common species is Sthenelais boa , easily told from 
other common scale worms by its elongated body and 100 or more 
pairs of scales. 

4. Dorsal surface convex, bristly; elongated setigerous dorsal 

lobes covering the dorsum 5 

4. Dorsal surface otherwise 6 

5. Notopodium a wide transverse ridge with 2 dorsal cirri and 
forked notosetae in transverse rows; with a dorsal crestlike 

"caruncle" on head (fig. 3) EUPHROSINIDAE (p. 67) 

With 6-7 branchiae on each notopodium, each with 1-4 branches: 
Euphrosine borealis . 

5. Notopodia with transverse membranous ridges supported by 
numerous spinelike setae (fig. 4); small, flattened, ellip- 

The only reported species is Spinther citrinus (Stimpson, 1854) , 

a small dredged form, lemon yellow, resembling in color the 
sponges with which it is associated. 

6. Anterior end bearing conspicuous filaments, or feathery ten- 
tacles, or conspicuous bristles, or golden setae (paleae) , so 

that prostomium is usually more or less concealed 7 

6. Anterior end with prostomium not concealed by such special 

outgrowths as may be present 12 

7. Anterior tentacles (branchial crown) are pinnate (feather- 
like) ; worms almost always form tubes 8 

7. Anterior end with flattened golden setae (paleae) forming 
an operculum (figs. 108, 110) ; tubes are of cemented sand 
grains, stiff and brittle 9 

7. Anterior end bearing many long filamentous (threadlike) out- 
growths; tubes of some are distictive, but lacking in ethers 10 

7. Anterior end somewhat concealed by a cage of long setae di- 
rected forward; body heavily papi Hated 


8. Tubes calcareous; one of tentacles modified to a pluglike 

stalked operculum SERPULIDAE (p. 77) 

8. Tubes flexible, leathery, parchment-like, sandy, or mucoid 

(except in the tiny Fabricia ) ; no operculum SABELLIDAE (p. 76) 

9. Tubes of sand, of regular conical form, open at both ends 

. „ ............ .PECTINARIIDAE (p. 74) 

The commonly taken species is Pectinaria gouldii (figs. 109, 
110) . 
9. Tubes of sand, convoluted and solidly cemented onto stones, 

shells, or into masses; common SABELLARIIDAE (p. 76) 

Polychaeta 61 

10. Filamentous outgrowths much concentrated on the head end; the 
body is divisible into two distinct regions ("thorax" and "ab- 
domen"); branching gills may be present in addition to filaments 11 

10. Filamentous outgrowths start back of the head (actual prostomium 
is bare) and are also borne on several to many segments of the 
body; never branching gills near anterior end; body not divis- 
ible into two distinct regions CIRRATULIDAE (p. 66) 

11. Anterior filaments not retractile into mouth, but tend to creep 
out radially if worms are undisturbed; prostomium seen only by 
lifting aside filaments; branchiae often present on dorsum be- 
hind filaments; well represented locally TEREBELLIDAE (p. 81) 

11. Anterior tentacles retractile into mouth, leaving pointed 
branchiae exposed and extending over anterior end; tapering 

at rear; not common in most collecting AMPHARETIDAE (p. 63) 

12. Anterior end with pair of long, coiling tentacular processes 
(often referred to as "palps") which may, however, be missing 

if specimen has been handled roughly 13 

12. Prostomium with obvious antennae (not minute relative to size 

of prostomium) ; with or without fleshy oral palps 15 

12. Prostomium with minute antennae; without fleshy oral palps 21 

12. Prostomium without antennae, or with a single median antenna; 

without tentacular cirri; or prostomium is indistinct 23 

13. Body segments similar in size and setae (except that the 5th 
may be much modified in Polydora ) ; body, although showing grad- 
ual transition, is not sharply divisible into distinct regions 

SPIONIDAE (p. 78) 

13. Body divisible into 2 or 3 distinct regions with dissimilar seg- 
ments 14 

14. Small, slender worms with spatulate prostomium; coiling ten- 
tacles bear a fringe of papillae or short branches on one side; 

no tube MAGELONIDAE (p. 69) 

14. Larger worms in distinctive leathery or chitinous tubes; para- 
podia are complexly lobed CHAETOPTERIDAE (p. 65) 

15. Dorsal cirri flattened, leaf like, often overlapping; prostomium 
with 2 eyes and 4-5 antennae; lacking oral palps; smallish, elong- 
ate worms of delicate build PHYLLODOCIDAE (p. 74) 

15. Dorsal cirri of typical tentacular or slender tapered form 16 

16. (With one rare exception) always with parapodial ligules (rounded, 
leaflike, or straplike dorsal lobes of parapodia) bearing dorsal 
cirri; prostomium with 4 eyes and 2 antennae; 2 biarticulated oral 

palps NEREIDAE (p. 70) 

16. Lacking parapodial ligules 17 

17. With 1-8 lateral pairs of tentacular cirri on peristomial seg- 
ment; small worms 18 

17. Without such peristomial cirri, or with a pair of short dorsal ten- 
tacles on nuchal region (fig. 16); dark complex jaw apparatus 19 

62 Polychaeta 

18. Prostomium with biarticulate oral palps; 2-3 antennae, 4 eyes, 

2-8 pairs of peristomial cirri HESIONIDAE (p. 68) 

Several species present locally, some of which are commensal. 
The most common species is the small, dark, short-bodied Podarke 
obscura (fig. 79) . 

18. Prostomium with 2 oral palps (sometimes reduced or fused), 3 
antennae, and 4-6 eyes; 1-2 pairs of peristomial cirri; the 
dorsal parapodial cirri often long, moniliform (beaded) , and 
conspicuous; a large and difficult family of small and beauti- 
ful worms SYLLIDAE (p. 79) 

19. Prostomium with 2 articulated antennae and 2 elongated palps; 

without branchiae DORVILLEIDAE (p. 66) 

19. Prostomium with 1-7 antennae, 2 low globular palps, best seen 

ventrally; branchiae simple, spiral, pectinate, or lacking 20 

20. With 5 occipital antennae (seem to be borne at rear of prosto- 
mium, but without frontal antennae; sometimes a pair of short 

nuchal tentacles (figs. 17-19) EUNICIDAE (p. 66) 

20. With 5 occipital antennae plus 2 frontal antennae on prostomium; 
occipital antennae with distinctly ringed bases (fig. 16) 

ONUPHIDAE (p. 71) 

21. Prostomium subquadrate, 1 pair of small pointed antennae at 
front corners, a second concealed at sides; parapodia biramous 
with lamellae well developed; body flattened, with distinctive 

"shimmying" motion NEPHTYIDAE (p. 70) 

21. Prostomium conical, long and tapering, with 4 minute antennae 
set in a cross at tip; parapodia small and lack lamellae; bodies 

pink, cylindrical, and pointed at both ends, coiling 22 

22. Segments bi- or triannulate; parapodia either uni- or biramous, 
but of one form for whole length of body; common 


22. Segments uni-annulate; parapodia of front part of body uniramous, 

biramous in rear GONIADIDAE (p. 68) 

23. Some of body segments much longer than wide 24 

23. Segments shorter than, or not much longer than, wide 25 

24. Segments remarkably long; anal segment with a funnel, or a 
flattened plate, or spatulate (spoon-like) ; prostomium hood-like 

or with flattened plate; "bamboo worms" MALDANIDAE (p. 69) 

24. Anal segment otherwise; prostomium with or without a slashed 

branchial membrane (fig. 107) OWENIIDAE (p. 73) 

25. Body very long and with small parapodia, somewhat resembling 
elongated earthworms; proboscis has dark jaw-pieces (not visible 

unless everted) ; 2 superficially very similar families 26 

25. Form various; proboscis without dark jaw-pieces 27 

26. Setae consisting of (1) simple, pointed, "limbate" setae 
(flattened back of tips) and (2) "hooded" setae (hooked 

setae with tips encased in a transparent drop of material) 


26. Setae limbate, with or without stout aciculum-like setae; 

without hooded setae ARABELLIDAE (p. 64) 

Polychaeta 63 

27. Parapodia of at least some segments bear typical uncinigerous tori 
(a torus is a simple raised area representing a low parapodial 
ramus; uncini are short embedded hooklike setae set in rows) 28 

27. Parapodia with capillary (simple, pointed) setae; typical 

uncinigerous tori absent, but palisaded rows of crochets (elong- 
■ate hooked setae) on low mounds may be present anteriorly 29 

27. Parapodia with filiform postsetal notopodial lobes, with strap- 
like branchiae dorsal to notopodia on some segments; prostomium 
subconical, with or without a median antenna; small, threadlike 
worms, living in mucous tubes in mud or sand 


28. Without branchiae, or when present, only in posterior region; 
body extremely long, slender, and fragile; characteristically 

purplish-red, found in mud CAPITELLIDAE (p. 65) 

28. With 11-13 pairs of dorsal branchiae in middle part of body; 

heavy-bodied; "lug worms" ARENICOLIDAE (p. 64) 

29. Prostomium bilobed or T-shaped; branchiae usually 4 pairs 
limited to anterior few segments, arborescent; integument 

checkered SCALIBREGMIDAE (p. 77) 

29. Prostomium pointed; branchiae are either numerous pairs or 

lacking altogether 30 

30. More or less short bodied, pointed at both ends; parapodia re- 
duced to bundles of setae, all simple, capillary; a ventral 
groove present in some; branchiae along sides of body or lack- 
ing „ OPHELIIDAE (p. 72) 

30. Elongate worms with pointed, actively inquiring heads; an- 
terior thoracic part of body often with flattened dorsum with 
parapodia lateral; posterior abdominal segments with parapodia 
dorsal; with paired branchiae dorsal to parapodia, the para- 
podia and branchiae covering the back like a furry coat 



1. Family Ampharetidae 
List of Species 

Amage auricula Malmgren, 1866. Dredged. 

Ampharete arctica Malmgren. 1866. Dredged. 

Ampharete acutif rons (Grube, 1850) . Dredged. 

Amphicteis qunneri (Sars, 1835). Dredged. 

Asabellides oculata (Webster, 1879). Dredged. 

Hypaniola gravi Pettibone, 1953. In shallow ponds of low salinity. Figure 5. 

Melinna cristata (Sars, 1851) . Dredged. 

Samytha sexcirrata (Sars, 1856) . Dredged. 

Samythella elongata Verrill, 1873. Dredged. 

2. Family Amphinomidae (not in key; rare) 
List of Species 

Amphinome rostrata (Pallas, 1766). Gulf Stream fauna, on floating weed, etc. 
Hipponoe gaudichaudi Audouin and Milne Edwards, 1830. On floating objects-, barnacles, 

etc . 
Paramphinome pulchella Sars, 1872. Dredged. 
Pareurythoe borealis (Sars, 1862) . Dredged. 

64 Polychaeta 

3. Family Aphroditidae 

1. Thick felty layer completely covers elytra; dredged in 

mud Aphrodita hastata Moore, 1905. 

1. Loose felty layer, not completely concealing elytra; 

dredged in sand Laetmatonice f ilicornis Kinberg, 1855. 

4. Family Arabellidae 

1. Parapodia without heavy projecting acicular setae; pro- 
stomium with four eyes in row on posterior margin (may 
be overlooked under fold of peristomium) (fig. 101) 
Arabella iricolor 

1. Parapodia provided with stout projecting aciculum-like 

setae (fig. 102) 2 

2. Prostomium conical, with four eyes in row on posterior 

margin Notocirrus spiniferus 

2. Prostomium spatulate, flattened dorsoventrally, without 

eyes Drilonereis 3 

3. Parapodia small, inconspicuous on anterior segments (fig. 
102); in far posterior region, parapodia distinctly bi- 
labiate (fig. 103) Drilonereis lonqa 

3. Parapodia prominent from first setiger on, similar through- 
out length of body Drilonereis magna 

List of Species 

Arabella iricolor (Montagu, 1804) . Formerly called A. opalina . Common in sandy 

Drilonereis lonqa Webster, 1879. Common in sandy mud. 

Drilonereis magna Webster and Benedict, 1887. Rare, dredged. 

Notocirrus spiniferus (Moore, 1906) . Low water and dredged. Young stages are 

thought to live parasit.ically in body cavity of Diopatra cuprea (Pettibone, M., 
1957. Endoparasitic polychaetous annelids of the family Arabellidae with de- 
scriptions of new species . Biol. Bull., 113 : 170-187). 

5. Family Arenicolidae 

1. 19 setigerous segments; 12-13 pairs of gills; found north of 

Cape Cod Arenicola marina 

1. 17 setigerous segments; 11 pairs of gills 2 

2. Firm, stout, dark blackish-green; commonly 15-30 cm long; 
castings formless; egg mass a gelatinous streamer up to a 

meter long Arenicola cristata 

2. Soft and limp; color pale pinkish-tan; length rarely exceeds 
15 cm; castings cylindrical, coiled; gelatinous egg mass firm 
and egg shaped Arenicola brasiliensis 

List of Species 
Arenicola b rasiliensis Nonato, 1958 (= A. caroledna Wells, 1961). This species 

Polychaeta 65 

prior to 1961 was not formally recognized as distinct from A. crista ta . al- 
though the difference had been noted years previously by the MBL's distin- 
guished former Collector, Mr. Milton "Sam" Gray. " Caroledna " is generally 
found in sand. 

Arenicola cristata Stimpson, 3 856. See above. This large Arenicola is the form 
traditionally used for dissection in the MBL Invertebrate Zoology Course. 
It prefers quieter waters and muddier sand than does A. brasiliensis , and 
is becoming increasingly scarce, in part because of dredging of harbors. 

Arenicola marina (Linnaeus, 1758). Occurs north of Cape Cod. 

6. Family Capitellidae 

1. Capillary (slender, pointed) setae on first 5 setigers; 

body very slender Heteromastus f iliformis 

1. Capillary setae on first 7 setigers Capitella capitata 

1. Capillary setae on first 11 setigers Notomastus 2 

2. Capillary setae in both noto- and neuropodia of first 
setiger; body blood-red anteriorly, light yellowish 

posteriorly Notomastus latericeus 

2. Setae only in notopodium of first setiger; body dark 
purplish -brown with bluish iridescence anteriorly 
Notomastus luridus 

Note: Capitellids, because of their extreme length and thinness, are diffi- 
cult to collect entire, and hence to identify. 

List of Species 

Capitella capitata (Fabricius, 1780). Common, in sandy mud. 
Heteromastus f iliformis (Claparede, 1864) . Common, in mud. 
Notomastus latereus Sars, 1851. In muddy sand. 
Notomastus luridus Verrill, 1873. In muddy sand. 

7. Family Chaetopteridae 

1. Tube U-shaped, parchment-like, with both ends open to sur- 
face; worm large, thick, well known 

Chaetopterus variopedatus 

1. Tube long, slender, transparent, annulated; buried ver- 
tically in sand; worm long and slender 
Spiochaetopterus oculatus 

List of Species 

Chaetopterus variopedatus (Renier, 1804) . Formerly C. perqamentaceus . Is fairly 
common, but not entirely dependable for experimental work because it may be- 
come very scarce in certain years. The distinctive tube may have led to over- 
collecting . 

Spiochaetopterus oculatus Webster, 1879. Uncommon; dredged in mud in Buzzards Bay 

and has been found by Mr. Gray on certain flats both north and south of the Cape. 

66 Polychaeta 

8. Family Cirratu lidae 

1. With 2 elongate grooved tentacles ("palps") arising from 
forward of the first setiger, in addition to the other 
filamentous outgrowths 3 

1. With numerous filaments , but without the differentiated 

grooved palps Cirratulus 

2. Prostomium bluntly conical with 2-9 eyes on each side in 
a transverse row or united in an arc (fig. 46) 

Cirratulus cJLrratus 

2. Prostomium conical, slightly acute; without eyes 

Cirratulus grandis 

3. Only 4-6 pairs of rather short filaments in addition to 
palps (fig. 45) ; dark, almost black; forms a calcareous 
tube mass; in calcareous shell or coral matrix 

Dodecaceria coralii 

3. With numerous filaments in addition to 2 tentacular palps 

(figs. 47, 48) 4 

4. With capillary (simple, pointed) setae exclusively in both 

rami of parapodia (fig. 47) Tharvx acutus 

4. With acicular setae or unidentate crochets in addition to 
capillary setae; crochets in rows almost encircling the 
body in posterior region (fig. 48) Chaetozone setosa 

List of Species 

Chaetozone setosa Malmgren, 1867. Dredged. 
Cirratulus cirratus (0. F. Muller, 1776) . Dredged. 
Cirratulus grandis Verrill, 1873. Common. 
Dodecaceria coralii (Leidy, 1855) . Dredged. 
Tharvx acutus Webster and Benedict, 1887. Dredged. 

9. Family Dorvilleidae 

1. Prostomium without eyes; in sand 

Stauronereis caecus (Webster and Benedict, 1884) 

1. Prostomium with 4 eyes (fig. 21) ; in sandy mud and in "weed" 
in Lagoon Pond (Martha's Vineyard) 

Stauronereis rudolphi (Delle Chiaje, 1828) 

10. Family Eunicidae 

1. With 2 short nuchal tentacles on the dorsal side of 

second segment (fig. 17) Eunice 3 

1. Without such nuchal tentacles Marphysa 2 

2. Prostomium bilobed (fig. 18); branchiae start on about seg- 
ment 20 (19-40) and continue nearly to end of body with 1-8 

filaments per branchia Marphysa sanguinea 

2. Prostomium rounded (fig. 19); branchiae start on segments 
12-15, relatively few in number (12-21 pairs) with 7-19 
filaments per branchia Marphysa bellii 

Polychaeta 67 

3. Acicula and subacicular hooks black; branchiae begin on 

setigers 7-9, and continue nearly to rear of body; tube 

papery Eunice norveqica 

3. Acicula and subacicular hooks yellow; branchiae begin on 

third setiger, and continue on to setigers 40-50, leaving 

middle and rear of body without branchiae; tubes thin, 

rough with debris Eunice pennata 

List of Species 

Eunice norveqica (Linnaeus, 1767). Includes E. f loridana Ehlers. Dredged. 
Eunice pennata (0. F. Muller, 1776) . Dredged. 

Marphysa bellii (Audouin and M. Edwards, 1833) . Rare, in muddy sand. 
Marphysa sanguinea (Montagu, 1815) . Common, in muddy sand. 

11. Family Euphrosinidae 
List of Species 

Euphrosine cirrata Sars, 1862. Dredged. 

Euphrosine borealis Oersted, 1843. Dredged; fig. 3. 

Euphrosine armadillo Sars, 1851. Dredged. 

12. Family Flabelligeridae 

1. Body covered with thick mucous mantle containing long, 
stalked papillae; neurosetae are compound hooks 
Flabelligera af finis 

1. Body without mucous mantle, covered with simple (not 

stalked) papillae; neurosetae are simple 2 

2. Both neuro- and notosetae are long, capillary; papillae 

numerous and remarkably long Diplocirrus hirsutus 

2. Neurosetae are stout simple hooks 3 

3. Hooked neurosetae begin on fourth setiger; body covered 
with elongate papillae, agglutinated with sand and mud 

Pherusa plumosa 

3. Hooked neurosetae begin on fifth setiger; body covered 

with short papillae (fig. 7) Pherusa affinis 

List of Species 

Brada granosa Stimpson, 1854. Dredged. 

Brada villosa (flathke, 1843) . Dredged. 

Diplocirrus hirsutus (Hansen, 1879) . Dredged. 

Flabelligera affinis Sars, 1829. Dredged. 

Pherusa affinis (Leidy, 1855) . Formerly Trophonia affinis ; in mud, shallow water, 

and has been taken at night light at WHOI dock. 
Pherusa plumosa (Muller, 1776). Dredged. 

13. Family Glyceridae 

1. Without branchiae; parapodia with single postsetal lobe 

and two unequal presetal lobes (fig. 89) 

Glycera capita ta 

1. Branchiae present (but may be retracted); parapodia with 

two presetal and two postsetal lobes (figs. 90, 91) 2 



Glycera dibranchiata 

Glycera americana 

Branchiae non-retractile at upper and lower edges of 
parapodia (fig. 90) 

Branchiae completely retractile, many lobed (fig. 91) , 
arising from grooves on posterior side of base of noto- 
podia (opening visible when branchia is retracted) 

List of Species 

Glycera americana Leidy, 1855. Common in muddy sand. 
Glycera capita ta Oersted, 1843. Dredged, in muddy sand. 
Glycera dibranchiata Ehlers, 1868. Common, in muddy sand. 

14. Family Goniadidae 


1. Presetal lobes of neuropodia bilobed (may be simple 

on some 18 of anterior segments) (figs. 92, 93) 2 

1. Presetal lobes of neuropodia simple, not bilobed; 
proboscis with about 28 chitinous, V-shaped, black 

chevrons Goniadella gracilis 

2. Proboscis without chevrons Ophioglycera gigantea 

2. Proboscis with 7-11 chevrons (fig. 88) 

Goniada maculata 

List of Species 

Goniada maculata Oersted, 1843. Dredged in mud. 

Goniadella gracilis (Verrill, 1873). Small, usually escapes notice; in fine sand. 

Ophioglycera gigantea Verrill, 1885. Rare, in mud flats. 

15. Family Hesionidae 
List of Species 

Gyptis vittata Webster and Benedict, 1887. Rare; under rocks. 

Microphthalmus aberrans (Webster and Benedict, 1887) . In sand; associated with 

terebellids Lysilla alba and Enoplobranchus sanguineus . 
Microphthalmus sczelkowii Mecznikow, 1865. In sand. 
Nereimyra punctata (0. F. Muller, 1776) . Dredged. 
Parahesione luteola (Webster, 1880). In sands, mud, commensal with Upogebia af - 

f inis . 
Podarke obscura Verrill, 1873. Common .in muddy sand and eel grass; may swarm 

at surface or to night light. Easily recognized by its very dark brown color. 

16. Family Lumbrineridae 

1. With palmately branched branchiae on some parapodia 

(figs. 96, 97) Ninoe nigripes 

1. Without branchiae Lumbrineris 2 

2. Prostomium extremely elongated, 2 to 3 times as long 

as wide (fig. 98) Lumbrineris acuta 

2. Prostomium short, conical (fig. 99) 3 

Polychaeta 69 

3. Body long, slender, small; acicula yellow; posterior 
parapodia with elongated finger like process (Fig. 100) 
Lumbrineris tenuis 

3. Body stouter; acicula black; parapodia without espec- 
ially elongated process Lumbrineris fragilis 

List of Species 

Lumbrineris acuta (Verrill, 1875). Rare. In muddy sand. 
Lumbrineris fragilis (0. F. Miiller, 1776) . Common in muddy sand. 
Lumbrineris tenuis (Verrill, 1873) . Common in muddy sand. 
Ninoe nigripes Verrill, 1873. Common. Dredged in mud. 

17. Family Magelonidae 

Magelona rosea Moore, 1907. Common in muddy sand. The only polychaete genus re- 
ported to have haemerythrin as a blood pigment. 

18. Family Maldanidae 

1. Anus within a funnel shaped structure (fig. 72) 2 

1. Anus associated with (dorsal to) an obligue structure 

(figs. 73, 74) 3 

2. With deep membranous collarette (fig. 71) on fourth 
setigerous segment; 18 setigerous segments; two color 
phases: pale with red nodes, or green (in mud) 

Clvmenella torguata 

2. Without collarette on fourth setiger; setigerous seg- 
ments variable in number (19-42) 

Clvmenella zonalis 

3. Anus dorsal to anal disc; anal segment forms a spatu- 
late lobe dorsally and funnel-like concavity ventrally 
(fig. 73) ; anterior end speckled with black or dark 

purple; mud tube • Maldanopsis elonqata 

3. Pygidium an obliquely truncated flat oval plate (fig. 74) 
with short lateral marginal incisions 
Maldane sarsi 

List of Species 

Asychis biceps (Sars, 1861) . Dredged. 
Axiothella catenata (Malmgren, 1856) . Dredged. 

Clymenel 1 a tnrqiiaha (Leidy, 1855). Very common in sand; forms sandy tubes. 
Clymenella zonalis (Verrill, 1874) . 
Leiochone dispar (Verrill, 1873) . Dredged. 

Maldane sarsi Malmgren, 1865. Common in very muddy sand. 
Maldanopsis elongata (Verrill, 1873) . Common in mud. 
Nicomache lumbricalis (Fabricius, 1780). Dredged. 
Petaloproctus tenuis (Theel, 1879) . Dredged. 

Praxillella gracilis (Sars, 1861). Dredged. One sp of Praxillella reported com- 
mon in mud beTow water at Hadley Harbor (Mr. Gray) . 
Praxillella praetermissa (Malmgren, 1866). Dredged. 
Praxillella ornata Verrill, 1880. Dredged. 
Rhodine loveni Malmgren, 1865. 

70 Polychaeta 

19. Family Nephytidae 

1. Branchiae (in form of a sickle shaped cirrus between 
dorsal and ventral ramus of each parapodium) curved 
inwards (fig. 77) ; not common Aglaophamus circinata 

1. Branchiae (see above) curved outwards (fig. 78) ; common 

Nephtys 2 

2. Tentacular segment with enlarged neuropodial lobe later- 
al to the setigerous lobe; dorsal tentacular cirri lack- 
ing on notopodium (fig. 76) 3 

2. Tentacular segment without enlarged neuropodial lobe; 
tiny dorsal tentacular cirri present on first notopodium 

(fig. 75) 4 

3. Ventral tentacular cirri anterior to widest part of 

enlarged tentacular segment Nephtys pi eta 

3. Ventral tentacular cirri lateral and continuous with 
widest part of enlarged tentacular segment (fig. 76) 

Nephtys bucera 

4. Both anterior and posterior parapodial lamellae about 
equally well developed, enclosing the conical acicular 

lobes ; setae dark Nephtys incisa 

4. Anterior parapodial lamellae rudimentary; posterior 

lamellae large, foliaceous; setae light Nephtys caeca 

List of Species 

Aglaophamus circinata (Verrill, 1874) . Dredged in mud. 
Nephtys bucera Ehlers, 1868. Common, in sand. 
Nephtys caeca (Fabricius, 1780) . Common, in muddy sand. 
Nephtys incisa Malmgren, 1865. Dredged. Common in mud. 
Nephtys picta Ehlers, 1868. Common, in muddy sand. 
Nephtys ciliata (0. F. Mullcr, 1789) . Dredged in mud. 

20. Family Nereidae 

1. Parapodia essentially uniramous , without ligules 

Lycastopsis pontica 

1. Parapodia biramous, with ligules (figs. 12-15) 2 

2. Paragnaths (denticles) (fig. 9) of proboscis small and 
pectinate (in comblike rows) ; peristomial cirri very long , 
to segments 10 or 15; often seen in heteronereid form at 

night lights Platynereis dumerilii mega lops 

2. Paragnaths of proboscis larger, conical (figs. 8, 10, 11) 3 

3. Notopodia of anterior parapodia with 2 ligules 4 

3. Notopodia of anterior parapodia with 3 ligules, the 

upper ligule (bearing cirrus) flattened 5 

4. Parapodial ligules sharply conical 

Nereis ( Nereis ) grayi 

4. Parapodial ligules short, thick, evenly rounded (fig. 13); 
worm may occur as a heteronereid 
Nereis ( Nereis ) pelagica 

Polychaeta 71 

5. Upper ligules of posterior parapodia elongated, strap- 
like, with cirrus near tip (fig. 12) ; often seen as the 
heteronereid at night light 
Nereis ( Neanthes) succinea 

5. Upper ligule large, broadly leaf-like, with cirrus in- 
serted near base (fig. 14) ; the most common large local 
nereid Nereis ( Neanthes ) virens 

5. Upper ligule triangular, with cirrus inserted near base 
(fig. 15) ; a small pale worm; very numerous tiny para- 
gnaths in oral ring (figs. 10, 11) 
Nereis ( Neanthes ) arenaceodonta 

List of Species 

Lycastopsis pontica (Bobretzky, 1872). Found in high intertidal. A primitive, 
aberrant nereid. 

Nereis ( Neanthes ) arenaceodonta Moore, 1903. Small; common in pilings and in al- 
gae; occasionally at night light. Includes N. caudata Delle Chiaje. 

N. ( Neanthes ) succinea (Frey and Leuckart, 1847) . Well known to embryologists 
as Nereis limbata . Common in brackish waters and in the peaty banks of es- 
tuaries. Usually taken as heteronereids at night light. 

N. ( Neanthes ) virens Sars, 1835. Very common in muddy sand and in more saline con- 
ditions than N. succinea . 

N. (Nereis ) grayi Pettibone, 1956. In mud, in tubes of Maldanopsis elongata . 

N. ( Nereis ) pelaqica Linnaeus, 1758. Common. On algae. 

Platynereis dumerilii mega lops (Verrill, 1874) . This form exhibits a unique 
copulation and internal fertilization well known to Woods Hole embryologists. 
It is regarded by Pettibone as a subspecies of P. dumerilii , but its method 
of reproduction is such that it is reproductively isolated from the exter- 
nally fertilizing typical P_. dumerilii of Europe. 

21. Family Onuphidae 

1. Peristomial segment without a pair of cirri; tube horny, 

transparent Hyalinoecia tubicola 

1. Peristomial segment with a pair of short tentacular 

cirri (fig. 16) 2 

2. Branchiae large and spiraled; worm large and wonder- 
fully iridescent; forms long stout vertical tube in 
sand, the projecting end cluttered with attached debris 
and shells 

Diopatra cuprea 

2. Branchiae not spiraled Ormphi .q 3 

3. Branchiae pectiniform or with 2 or more filaments where 

best developed subgenus Onuphis 5 

3. Branchiae simple, cirriform subgenus Nothria 4 

4. Branchiae begin on segments 10-13; parapodia of first 
segment greatly enlarged; tube free, parchment-like, 
flattened, covered with pebbles and shells 

Onuphis ( Nothria ) conchy lega 

4. Branchia begin on first segment; parapodia of first 
segment not enlarged; muddy tube 
Onuphis ( Nothria ) opalina 

72 Polychaeta 

5. Branchiae begin on segments 5-6 , with a maximum of 
4-5 pectiniform filaments 

Onuphis ( Onuphis ) quadricuspis 

5. Branchiae begin on first segment, with a maximum of 
5-7 pectiniform filaments 
Onuphis ( Onuphis ) eremita 

List of Species 

Diopatra cuprea (Bosc, 1802). The prominent tubes, set in sand, are common and 

easily recognized. 
Hyalinoecia tubicola (0. F. Miiller, 1776). Dredged in mud. 
Onuphis ( Nothria ) conchyleqa Sars, 1835. Dredged in mud; tube free, scabbard 

Onuphis ( Nothria ) opalina (Verrill, 1873) . Dredged in mud; tube of mud. 
Onuphis ( Onuphis ) eremita Audouin and M. Edwards, 1833. Dredged in sand and mud; 

tube of sand or mud. 
Onuphis ( Onuphis ) quadricuspis Sars, 1872. Dredged; tube of mud. 

22. Family Opheliidae 


1. Body stout, grub like; no ventral groove 

Travisia carnea 

1. Body Amphioxus shaped, with ventral groove for whole 

length Ammotrypane aulogaster 

1. Ventral groove only behind segments 10-12 

Ophelia 2 

2. Branchiae 18 pairs, crenulate on sides 

Ophelia denticulata 

2. Branchiae 11-15 pairs, smooth Ophelia bicornis 

List of Species 

Ammotrypane aulogaster Rathke, 1843. Dredged. 

Ophelia bicornis Savigny, 1818. Only north of Cape. 

Ophelia denticulata Verrill, 1875. 

Travisia carnea Verrill, 1873. On sand flats, Naushon Island (Mr. Gray). 

23. Family Orbiniidae 


1. Thoracic neuropodia with numerous postsetal papillae in ver- 
tical rows (fig. 104) ; branchiae begin on fifth seti- 
ger; large and robust worms Orbinia ornata 

1. Thoracic neuropodia without rows of papillae (1-3 
postsetal papillae present in some) ; branchiae start 

on setigers 9-32 Scolopios 2 

2. With a cirrus between the dorsal and ventral para- 
podial rami of anterior abdominal parapodia (figs. 

105, 106) 3 

2. Without an interramal cirrus Scolopios acutus 



3. With ventral papillae on some anterior segments 
(up to 9 in a row on each side of segments 14- 
24) Scoloplos riseri 

3. Without ventral papillae 4 

4. With 2-3 subpodial papillae in transitional region 
between thoracic and abdominal regions, followed 
by an entire flattened subpodial flange (may be 
undulate but not incised; fig. 105) 

Scoloplos robustus 

4. With 2 subpodial papillae in transitional region, 
continuing as 2 subpodial lobes (thus a notched or 
incised subpodial lobe; fig. 106) 

Scoloplos fragilis 

List of Species 

Naineris quadricuspida (Fabricius, 1780). Dredged in mud. 

Orbinia ornata (Verrill, 1873) . Formerly called Aricia ornata , common in sandy 

shoals . 
Scoloplos acutus (Verrill, 1873) . Dredged in soft mud. 
Scoloplos fragilis (Verrill, 1873) . 
Scoloplos riseri Pettibone, 1957. 
Scoloplos robustus (Verrill, 1873). Common. In vertical burrows in muddy sand; 

has been called Haploscoloplos robustus . 

24. Family Oweniidae 


1. Prostomium rounded, without appendages; dredged 

Myriochele heeri Malmgren, 1867- 

1. Prostomium with a slashed branchial membrane (fig. 
107); tube distinctive, with a dense coating 
of flattened sand grains and bits of shell set 
in overlapping fashion like shingles; dredged 

Owenia fusiformis Delle Chiaje, 1844. 

25. Family Paraonidae 


1. Without dorsal antenna Paraonis 2 

1. With dorsal median antenna (fig. 80); branchiae 
begin on fourth segment 


2. Branchiae begin on segments 6-7, 9-14 pairs 

Paraonis gracilis 

2. Branchiae begin on fourth segment, 16-25 pairs 

Paraonis fulgens 

3. Median antenna long, filiform, extending to about 

segments 4-6; branchiae 9-10 pairs Aricidea quadrilobata 

3. Median antenna short (fig. 80), extending to first 
segment; branchiae 11-18 pairs 
Aricidea jef freysii 

74 Polychaeta 

List of Species 

Aricidea jef freysii (Mcintosh, 1879) . Dredged in mud. 

Aricidea quadrilobata Webster and Benedict , 1887. Dredged in soft mud. 

Paraonis fulgens (Levinsen, 1883). In sandy beaches. 

Paraonis gracilis (Tauber, 1879) . Dredged in soft mud. 

26. Family Pectinariidae 

List of Species 

Pectinaria ( Cystenides ) gouldii (Verrill, 1873) (fig. 108) . Intertidal; common. 
This is often referred to as Cystenides gouldii ; Cystenides is a subgenus of 
Pectinaria . Conical sand tube (fig. 109) is distinctive of family. 

Pectinaria granulata (Linnaeus, 1767). Dredged. 

Pectinaria hyperborea (Malmgren, 1866) . Dredged. 

27. Family Phyllodocidae 


1. Two pairs (one pair on each side) of peristomial cirri; 
prostomium triangular with 4 small antennae (fig. 56) 
Eteone 2 

1. Four pairs (2 pairs on each side) of peristomial tenta- 
cular cirri (figs. 60, 61) 3 

2. Elongate, slender; pale or yellow; dorsal cirri of 
middle region on body asymmetrical 

Eteone lactea 

2. Robust, dark green with light green transverse bands 
between segments; dorsal cirri of middle region sym- 
metrical, almost triangular Eteone longa 

3. Four prostomial antennae (figs. 57, 61) 4 

3. Five prostomial antennae (figs. 58, 59, 60) 6 

4. Prostomium cordiform (heart shaped) , with an occipital 
tubercle in the posterior notch (fig. 57) 

Phyllodoce ( Anaitides ) 5 

4. Prostomium oval, without an occipital tubercle (fig. 61) 

Paranaitis speciosa 

5. Ventral cirri oval, blunt, or only slightly pointed dis- 
tally, with a pointed ventral projection; body heavily 

pigmented Phyllodoce groenlandica 

5. Ventral cirri acutely pointed distally; green, banded 
with brown, with white marks on dorsum; dorsal cirri 

spotted Phyllodoce mucosa 

6. With 4-5 paired nuchal lappets posterior to prostomium; 
parapodia biramous; dorsal cirri plate-like, imbricated, 
resembling elytra of scale worms (fig. 58) 

Notophyllum americanum 

6. Without nuchal lappets; parapodia uniramous 7 

7. Three tentacular segments distinct dorsally (fig. 59) 

Eulalia 8 

7. First tentacular segment not distinct dorsally (fig. 
60) ; dorsal cirri cordiform; body greenish and yel- 
lowish, banded with grayish-green, reddish- brown, or 
light brown Eumida sanguinea 

Polychaeta 75 

8. Dorsal cirri elongate-lanceolate; greenish; body 

rather short Eulalia viridis 

8. Dorsal cirri thick, oval-obtuse; greenish with 

darker longitudinal lateral bands; very slender, 

elongate Eulalia bilineata 

List of Species 

Eteone lactea Claparede, 1868. Common in sandy mud. Includes E. alba Webster. 

Eteone lonqa (Fabricius, 1780). Includes E. robusta Verrill. 

Eulalia bilineata (Johnston, 1840) . In algae. May form epitokous sexual form. 

Eulalia viridis (Linnaeus, 1767). Common, on pilings. 

Eumida sanguinea (Oersted, 1843). Common, on pilings. 

Notophyllum americanum Verrill, 1885. Rare. Dredged. 

Paranaitis speciosa (Webster, 1880) . Rare. In sand. 

Phyllodoce ( Anaitides ) arenae W ebster, 1879. In muddy sand. May swarm at surface 

of water. 
Phyllodoce ( Anaitides ) qroenlandica Oersted, 1842. Dredged. 

Phyllodoce ( Anaitides ) maculata (Linnaeus, 1767). Common among rocks and algae. 
P hyllodoce ( Anaitides ) mucosa Oersted, 1843. Common in muddy sand. 

28. Family Polynoidae 

1. Elytra 12 pairs (fig. 1) 2 

1. Elytra, more than 12 pairs 3 

2. Elytral tubercles small, widely spaced; commensal in 
shells occupied by Pagurus pnllicaris 

Lepidonotus sublevis 

2. Elytral tubercles larger, crowded; very common, 

free living Lepidonotus squamatus 

3. 15 pairs of scales 4 

3. 40-50 pairs of scales; commensal in tubes of Amphi - 

trite ornata Lepidametria commensalis 

4. Anterior pair of eyes antero-ventral on prostomium, 
not visible dorsally (fig. 2) 

Harmothoe imbricata 

4. Anterior pair of eyes antero-dorsal on prostomium, 

visible dorsally Harmothoe extenuata 

List of Species 

Alentiana aurantiaca (Verrill, 1885). Dredged; commensal with anemone, Bolocera 

tuediae . 
Antinoella angusta (Verrill, 1874) . Dredged in mud. 
Arcteobia anticostiensis (Mcintosh, 1874) . Dredged in mud. 
Enipo gracilis Verrill, 1874. Dredged in mud; commensal with maldanid, Nicomache 

lumbricalis . 
Gattyana amondseni (Malmgren, 1867) . Dredged in mud. 
Gattyana cirrosa (Pallas, 1766) . Dredged in mud. 
Harmothoe ( Eunoe ) nodosa (Sars, 1860) . Dredged. 
Harmothoe ( Eunoe ) oerstedi (Malmgren, 1865) . Dredged. 
Harmothoe (Eunoe) spinulosa (Verrill, 1870) . Dredged. 

-jr Polychaeta 

Harmothoe ( Lagisca ) extenuata (Grube, 1840). Common; under stones, algae. 

Harmothoe imbricata (Linnaeus, r?67)Very common. This is the species usually en- 
countered; under stones, algae. 

Harmothoe fraqilis Moore, 1910. Dredged. 

Lepidametria commensalis Webster, 1879. Commensal in tubes of Amphi trite ornata . 

Lepidonotus squamatus (Linnaeus, 1758). Very common. Under stones. 

Lepidonotus sublevis Verrill, 1873. Dredged. Commensal in snail shells occu- 
pied by hermit crab Paqurus pollcaris . 

29. Family Sabellariidae 

Sabellaria vulgaris Verrill, 1873. Common. Dredged; on shells or in bryozoan 
nodules (fig. 110) . 

30. Family Sabellidae 

1. Tiny, of 10-12 setigerous segments; eyes on first segment 
and on rear of body (fig. 50) ; poorly defined muddy tubes, 
which worms may leave and move about, tail first 
Fabricia sabella 

1. Larger, of more than 12 setigerous segments 2 

2. Branchial filaments united by membrane for most of 
length; tube of mucus, transparent and thick 

Myxicola infundibulum 

2. Branchial filaments united by membrane for at least 
half their length; tubes inconspicuous, usually 
buried in sand or mud 3 

2. Branchial filaments often with eyes and united only 
at bases; tubes obvious, leathery, usually coated 

with sand 4 

3. Large ventral groove with flared sides on about 9 
posterior segments; collarette bilobed: notched 
mid-dorsally and with a small ventral slit 

Euchone rubrocincta 

3. Without ventral groove; collarette entire ventrally, 
with only a small mid-dorsal slit (fig. 51) 

Chone infundibuliformis 

4. Branchial filaments without eyes 

Potamilla neglecta 

4. Branchial filaments with eyes 5 

5. Eyes in pairs, 2-6 on each branchial filament, situa- 
ted in the transverse color bands; collarette 4-lobed 
by a small lateral notch on each side in addition to 
the dorsal and ventral slits (fig. 52) 

Sabella crassicornis 

5. Eyes not paired 6 

6. Branchial filaments with 2 irregular rows of many 
eyespots ; collarette bilobed by ventral and dorsal 
slits; body short and thick (figs. 53, 54) 

Sabella microphthalma 

6. Branchial filaments with 1-8 large "compound eyes" 
in a single row; collarette 4-lobed; body long and 
thin (fig. 55) Potamilla reniformis 

Polychaeta 77 

List of Species 

Chone infundibuliformis . Kr6"yer, 1856. 

Euchone rubrocincta (Sars, 1861) . 

Fabricia sabella (Ehrenberg, 1837) . This is the only one of our sabellids that can 

leave its tube and move about. 
Myxicola infundibulum (Renier, 1804). Dredged; noted for its giant axon. 
P otamilla neglecta (Sars, 1851). Dredged. 

Potamilla renif ormis (Linnaeus, 1788). Common; dredged on shells. 
Sabella crassicornis Sars, 1851. Dredged. 
Sabella microphthalma Verrill, 1873. 

31. Family Scalibregmidae 

1. Prostomium T-shaped, with frontal horns (fig. 20); body inflated 

anteriorly. In mud; intertidal and dredged. . Scalibregma inf latum Rathke, 1843. 

1. Prostomium bilobed; without frontal horns; body short and fusi- 
form Polyphysia crassa (Oersted, 1843) . 

32. Family Serpulidae 

1. Minute, flat coiled tubes on stones, algae. Spirorbis 2 

1. Irregular tubes up to several inches long, on rocks and shells 5 

2. Tubes coiled "dextrally" (figs. 62, 63, 64) 3 

2. Tubes coiled 'sinistrally" (figs. 65, 67) 4 

3. Tube white, opaque, shiny, smooth (figs. 62, 63) 

Spirorbis ( Dexiospira ) spirillum 

3. Tube white, opaque, thick and tough with 3 rounded longitudinal 
ridges produced into blunt teeth at opening (fig. 64); lined with 

dark layer Spirorbis ( Paradexiospira ) violaceus 

4. Tube without luster, chalky, without longitudinal ridges (fig. 65); 

eggs incubated in tube Spirorbis ( Laeospira ) bo realis 

4. Tube with 2-3 longitudinal ridges to almost smooth (fig. 67); 
eggs incubated in a brood pouch in operculum (fig. 68) 

Spirorbis ( Laeospira ) granulatus 

5. Operculum a two-tiered structure on a smooth peduncle (fig. 70); 
common; tube usually attached for most of length 

Hydroides ( Eupomatus ) dianthus 

5. Without an operculum; tube large white, cylindrical; erect and 

almost straight in its unattached distal portion Protula tubularia 

5. Two opercula, each a thin shallow membraneous cup with barbules 
on peduncle (fig. 69); tubes very slender (ca. 1 mm) in an inter- 
twining mass F iloqrana imp lexa 

List of Species 

F iloqrana i mplexa Berkeley, 1828. Dredged. 

Hydroides ( Eupomatus ) dianth us (Verrill, 1873) . This has been called Hydroi des hex- 

agonas , Serpula dianthus , Eupomatus dianthus , and Hydroides uncina ta . 
Protula tubularia (Montagu, 1803) . Dredged. 
Spirorbis ( Dexiospira ) spirillum (Linnaeus, 1758). Common. 
Spiro r bis ( Laeospira ) borealis Daudin, 1800. Common. 
Spirorb is ( Laeospira ) granulata (Linnaeus, 1767). Dredged. 
Spirorbis (P aradexiospira ) violaceus Levinsen, 1883. Dredged. 

78 Polychaeta 

33. Family Sigalionidae 
List of Species 

Leanira hystricus Ehlers, 1875. Dredged in mud. 

Leanira tetragona (Oersted, 1845) . Dredged in mud. 

Pholoe minuta (Fabricius, 1780). Dredged. 

Sigalion arenicola Verrill. 1879. Rare, in sand. 

Sthenelais boa (Johnston, 1873). Common, in Zostera root masses in certain areas, 

e.g., Lagoon Pond. 
Sthenelais limicola (Ehlers, 1864) . Dredged in sand and mud. 

34. Family Spionidae 

1. Fifth setiger very modified, with large dorsal setae 

(fig. 6) i Polydora 2 

1. Fifth setiger like the rest, unmodified 4 

2. Branchiae begin on setiger 6, continuing nearly to 
posterior end of body; color orange; pygidium with 
many papillae; bores in shells used by hermit crabs 

Polydora commensalis 

2. Branchiae begin on setiger 7, continuing nearly to 
posterior end; pygidium with an anal cup notched 

dorsally 3 

3. Prostomium clearly bifid anteriorly, and with a median 

occipital antenna (fig. 81) Polydora liqni 

3. Prostomium indistinctly bifid anteriorly; without a 

median occipital antenna Polydora ciliata 

4. Without branchiae; prostomium T-shaped (fig. 82) 

Spiophanes bombyx 

4. With 1 pair of branchiae dorsal to the 2 coiling palps 

Streblcspio benedicti 

4. With 4-5 pairs of branchiae, some pinnate, others 

smooth Prionospio spp. 

4. With numerous branchiae 5 

5. Branchiae begin on or about 13th setiger (11-20); 

about 14 pairs (7-28) Pyqospio eleqans 

5. Branchiae begin on first setiger, absent from last 
half or third of body; in brackish water (fig. 83) 

Scolecolepides viridi s 

5. Branchiae begin on second setiger, continue to rear 
of body; in sandy beaches (fig. 85) 
Scolelepis squama ta 

5. Branchiae begin on first setiger, continue to rear 

of body 6 

6. Prostomium conical; eyes nearly in a transyerse line 

(fig. 84) Dispio uncinata 

6. Prostomium inflated anteriorly; eyes set in a square 

(fig. 86) Spio 7 

7. Ventral lamellae greatly reduced; about 16 hooded cro- 
chets in each neuropodium; body dull green with red 
branchiae held erect over dorsum; common in thick fragile 

sand tubes on beaches Spio setosa 

7. Ventral lamellae little reduced; about 6 hooded crochets 

in each neuropodium (fig. 87) Spio f ilicornis 

Polychaeta 79 

List of Species 

Dispio uncinata Hartman, 1951. Burrows in sand. 

Laonice cirrata (Sars, 1851). Dredged. 

Polydora ciliata (Johnston, 1838). In shells. 

Polydora commensalis Andrews, 1891. A bright orange-red worm, commensal with her- 
mit crabs in snail shells; penetrates columnella of shell and not seen with- 
out breaking shell. Orange eggs attached in clusters in burrow. 

Polydora liani Webster, 1879. Makes vertical burrows in stiff mud or clay, often 
in estuaries. Eggs laid in capsules in burrows. 

Prionospio heterobranchia Moore, 1907. At low water mark and dredged, in mud. 

Prionospio steenstrupi Malmgren, 1867. Dredged in mud. 

Scolecolepides viridis (Verrill, 1873) . Penetrates further up estuaries than any 
other local polychaete. 

Scolelepis squamata (0. F. MiJller, 1789) . Includes Nerinides aqilis (Verrill, 
1873). Makes vertical tubes in sandy beaches. 

Spio f ilicornis (0. F. Miiller, 1776) . Dredged. 

Spio setosa Verrill, 1873. The fragile chimney-like sandy tubes are distinctive 
in beach areas. Common. 

Spiophanes b ombyx (Clax^arede, 1870) . Slender, branched, sandy tube. 

Streblospio benedicti Webster, 1879. Characteristically in estuaries. 

35. Family Sternaspidae (not in key) 

Sternaspis scutata (Renier, 1807). Distinctive, gray, grub- like worms, with a 
pair of horny ventral plates. Dredged in soft bottoms. 

36. Family Syllidae 

1. Without ventral cirri; palps reduced or lacking; often 
collected at night lights as sexually dimorphic males 
( " Polybostrichus ") and females ( "S acconereis " ) , the 
latter bearing sacs of white or brilliantly colored 
eggs Subfamily AUTOLYTINAE 4 

1. With ventral cirri; palps better developed 2 

2. Antennae and cirri moniliform (beaded); oral palps free 

or fused at base only Subfamily SYLLINAE 5 

2. Antenna and dorsal cirri smooth or indistinctly beaded 3 

3. Palps not fused, or fused only at base 

Subfamily EUSYLLINAE 7 

3. Palps fused for entire length; eggs and larvae attached 
along dorsal or ventral surfaces of the female 

Subfamily EXOGONINAE 11 

Subfamily Autolytinae 

4. Without long capillary swimming setae: the "stem" or 
asexually reproducing form of Autolytus (fig. 37) ; see key in 
Pettibone (1963) . 

4. With long capillary swimming setae; in "heterosyllid" 

form with body in 2 or 3 distinct regions; sexually di- 
morphic; both males and the females, carrying 1-3 sacs 
of eggs, are often taken at night-lights. The specific 
identification of the tiny worms is difficult; consult 
the more complete treatment by Pettibone (1963). 

80 Polychaeta 

Subfamily Syllinae 

5. Parapodia with simple setae only, few (2-5) in number, 

with bifid tips; dorsal cirri with - 60 articles Syllis spongiphila 

5. At least some of parapodia have compound setae 6 

6. Dorsal cirri alternately longer and shorter, with 20-40 
articles; blades of upper compound setae fringed, appear- 
ing bifid Syllis cornuta 

6. Dorsal cirri with 7-16 articles (figs. 23, 24); blades 
of compound setae do not appear bifid (fig. 25), but 
there may be heavy bifurcated simple setae (fig. 26) in 
some of the median segments Syllis gracilis 

Subfamily Eusyllinae 

7. With occipital or nuchal epaulettes; body short, of few 
segments; proboscis long, sinuous, armed with a complete 
circle of bicuspid teeth Amblyosyllis finmarchica 

7. Without occipital or nuchal epaulettes; proboscis straight 


8. Occipital flap or nuchal hood covers rear of prostomium 
(fig. 28); proboscis armed with semicircle of 6-7 large 

recurved teeth (fig. 29) Odontosyllis fulgurans 

8. Without nuchal hood 9 

9. Proboscis armed with a large anterior tooth and with 
numerous denticles (fig. 32); prostomium with 6 eyes 

(fig. 31) Eusyllis lamelligera 

9. Proboscis unarmed 10 

10. Antennae and dorsal cirri cylindrical; acicula large, 

knobbed (fig. 30) Streptosyllis sp. 

10. Antennae and dorsal cirri swollen, club shaped; with- 
out large knobbed acicula Syllides spp. 

Subfamily Exogoninae 

11. Two pairs of tentacular {peristomial) cirri ; 3 pairs of 
eyes; antennae and dorsal cirri fusiform (fig. 33) 

Brania clavata 

11. One pair of tentacular cirri 12 

12. Tentacular cirri 1 pair, similar to antennae; dorsal 
cirri swollen at base, tapering to narrow tip (fig. 36) 

Sphaerosyllis erinaceus 

12. Tentacular cirri 1 pair, rudimentary ; dorsal and ven- 
tral cirri cylindrical or club shaped, not swollen at 
base, median antenna fusiform, reaching nearly to end 
of palps (fig. 34) Exoqone 13 



13. Two pairs of eyes (fig. 35) Exogone dispar 

13. Three pairs of eyes (fig. 34) Exogone hebes 

Subfamily Autolytinae 
List of Species 

Autolytus alexandri Mal'!>gren, 1867. On algae. 
Autolytus cornutus A. Agassiz, 1863. Common on algae. 
Autolytus emertoni Verrill, 1881. Taken in plankton. 
Autolytus fasciatus (Bosc, 1802). Includes A. ornatus Verrill. 
Autolytus prismaticus (Fabricius, 1780). Dredged; in sponges. 
Autolytus prolifer (0. F. Muller, 1788). Common on pilings. 

Common on pilings. 

Subfamily Syllinae 
List of Species 

Syllis cornuta Rathke, 1843. Dredged; in sponges. 

Syllis gracilis Grube, 1840. Common, under stones or algae. 

Syllis spongiphila Verrill, 1885. Dredged; in muddy sand. 

Subfamily Eusyllinae 
List of Species 

Amblyosyllis f inmarchica (Malngren, 1867) . Includes Pterosyllis cincinnata Verrill) 

Eusyllis blomstrandi Malmgren, 1867. Dredged. 

Eusyllis lamelligera Marion and Bobretsky. Includes E. fragilis Webster. Common; 
dredged among shells, bryozoans. 

Odontosyllis fulgurans Claparede, 1864 . Common on pilings. Reproduces by lumin- 
escent swarmers at surface. 

Streptosyllis sp. 

Syllides longocirrata Oersted, 1845. Includes S_. convoluta Webster and Benedict. 
In sand. 

Syllides setosa Verrill, 1882. In mussel beds. 

Subfamily Exogoninae 
List of Species 

Brania clavata (Claparede, 1863) . Common, in mud, mussel beds. 

Brania wellfleetensis Pettibone, 1956. 

Exogone dispar (Webster, 1879) . Common, in mussel beds. 

Exogone hebes (Webster and Benedict, 1884) . 

Exogone veruqera (Claparede, 1868) . 

Parapionosyllis longicirrata (Webster and Benedict, 1884) . Common in muddy sand. 

Sphaerosyllis erinaceus Claparede, 1863. In sand or on pilings. 

37. Family Terebellidae 

With dorsally placed gills just behind tentacular fila- 
ments (usually contrasting with filaments in color) ; 
body usually with pronounced thickening of "thoracic" 


Lacking dorsal gills (branchiae) on anterior part of body; 
bodies less stout, and departing more from the fully ex- 
pressed "typical" terebellid form than the above group . . 


82 Polychaeta 

2. Gills arborescent or branching in tree-like fashion (fig. 

38) 3 

2. Gills consisting of one to many unbranched filaments 

(fig. 39) 8 

3. Three pairs of branching branchiae (note: one member of 

a pair may be very small or lacking) 4 

3. Two pairs of branching branchiae (note: as above) 6 

3. One pair of branching branchiae; numerous eyespots (fig. 

41) ; 16 thoracic setigerous segments 
Pista maculata 

3. With a single large branchia formed of 4 branchiae fused 
into one large trunk bearing 4 pectinate lobes (fig. 43) ; 

18 thoracic setigers Terebellides stroemi 

4. Setae on 40-50 segments; a large and common worm 

Amphi trite ornata 

4. Setae on 23-45 segments Amphi trite Johns toni 

4. Setae extend to posterior end of body 

Terebella lapidaria 

4. Setae on 17 segments 5 

5. Buccal segment with large lateral lobes, joined ventrally; 
the first of the three pairs of branchiae with long main 
stems, much longer than the last two pairs 

Loimia medusa 

5. Without enlarged lateral lobes on buccal segment; branchiae 

subequal Amphi trite af finis 

6. Setae on 17 segments (note: first setigerous segment 
lacks the neuropodial uncini) ; with lateral lobes on 

anterior segments 7 

6. Setae on 15 segments; numerous eyespots (fig. 40); with- 
out lateral lobes on anterior segments 

Nicolea venustula 

Note: The young of Nicolea are commonly seen as 'hydra-worms" in washings of 
Fiimis : they resemble creeping brown hydras with eyespots on cephalic ridge. 

7. Branchiae spirally branched, making (when contracted) a 
compact oval red pompom on a stalk (fig. 42); one branchia 
usually much larger than rest 

Pista crista ta 

7. Branchiae arborescent, not spirally branched, with large 

main trunk, usually unequal in size Pista palmata 

8. With 3 pairs of branchiae, each consisting of one simple 
long filament; numerous eyespots; 15 thoracic setigers 

Trichobranchus qlacialis 

8. With 2-3 pairs of branchiae, each made up of transverse 

rows of unbranched filaments (fig. 39) 9 

9. Two pairs of branchiae; numerous eyespots; notosetae be- 
gin on second gill bearing segment and continue to post- 
erior end Thelepus cincinnatus 

9. Three pairs of branchiae; no eyespots 10 

10. Notosetae begin on third branchial (gill bearing) seg- 
ment, continue on 17 segments; uncini begin on second 
setiger; tube membranous, encrusted Amphi trite cirrata 

10. Notosetae begin on first branchial segment and continue 
over a large part of body; uncini begin on fourth seti- 
ger; tube hard, coiled Streblosoma spiralis 

Polychaeta 83 

11. Very soft and fragile; transparent to yellowish; no 

setae; somewhat resembles a damaged Leptosvnapta when 

collected, but tentacles are terebellid-like Lysilla alba 

11. Fragile, elongate, blood-red body; setae on all seg- 
ments, but no uncini; branching red parapodia in mid 
body region: these look like gills, but note setae! 
(fig. 44) Enoplobranchus sanguineus 

11. Small worms, enveloped in their tentacles, in which 
red corpuscles circulate, but lack actual branchiae; 

setae and uncini present in part of body Polycirrus 12 

12. Red; setae on 18-25 segments; common Polycirrus eximius 

12. Red; setae on 11-13 segments Polycirrus medusa 

12. Lemon-yellow; setae on 24-32 segments Polycirrus phosphoreus 

List of Species 

Amphitrite af finis Malmgren, 1866. Dredged. 

Amphi trite cirrata 0. F. Miiller, 1771. Dredged. 

Amphitrite Johns toni Malmgren, 1866. Dredged. 

Amphitrite ornata (Leidy, 1855). Common. Forms muddy mounds on intertidal flats. 

Enoplobranchus sanguineus (Verrill, 1873). In mud; easily recognized by the bril- 
liant color. 

Loimia medusa (Savigny, 1818) . Dredged. 

Lysilla alba W ebster , 1879. In guite muddy sand, where it makes a surface depres- 
sion much like that of Leptosynapta . 

Nicolea venustula (Montagu, 1818). Usually seen as the tiny (immature) "hydra 
worms" in washings from algae. 

Pista cristata (0. F. Muller, 1776) . Occurs in mud, but tube is very rough and en- 
crusted with coarse pebbles. Common. 

Pista maculata (Dalyell, 1853). Dredged. 

Pista palmata (Verrill, 1873) . 

Polycirrus eximius (Leidy, 1855) . The commonly taken form. Red cells circulate 
in tentacles. 

Polycirrus medusa Grube, 1850. Dredged. 

Polycirrus phosphoreus Verrill, 1880. 

Streblosoma spiralis (Verrill, 1874) . Dredged. 

Terebella lapidaria (Linnaeus, 1767). 

Terebellides stroemi Sars, 1835. Dredged. 

Thelepus cincinnatus (Fabricius, 1780). Dredged. 

Tr i chobr anchu s glacialis Malmgren, 1866. Dredged. 


Fauvel, P., 1923. Polychetes E rrantes . Faune de France, Paris, _5: 1-488. 

Fauvel, P., 1927. Polychetes Sedentaires . Faune de France, Paris, j^: 1-494. 

Pettibone, M. H., 1963 . Marine Polychaete Worms of the New England Region, Part 1, 
Families Aphroditidae through Trochochaetidae . Bull. U. S. Nat. Mus., 227 : 

Pettibone, M.H., Ibid . Part 2, in preparation. 

Hartman, O., 1951. Literature of the Polychaetous Annelids. Vol. 1. Biblio- 

Hartman, 0., 1959. Catalogue of the polychaetous annelids of the world. Allan 
Hancock Foundation Publ., Occas . Paper 2_3: 1-628. (For additional bibliography 
and synonymy) . 

84 Chapter X 


Subphylum Chelicerata 

In the great group of chelicerates are included a variety of "arachnoid" types , most 
of which are in highly specialized terrestrial groups. Marine representatives include 
the primitive and ancient Xiphosur J da ("horseshoe crabe") , the small and specialized or- 
der of Pycnogonida ("sea spiders") of obscure affinities, and the marine mites (Order 
Acarina, Family Halacaridae) . Of the latter, about 20 species have been recorded from 
southern New England, but the study of these small creatures presents difficulties, and 
the interested student is referred to the comprehensive work of Newell, I. M. , 1947. 
A systematic and ecological study of the Halacaridae of eastern North America. Bull. 
Bingham Oceanogr. Coll., 1J3: 1-232. 

Class Xiphosurida 

Limulus Polyphemus (Linnaeus), the common "horseshoe crab", has been almost univer- 
sally known as Limulus , except for a brief period in which the name Xiphosura polyphe - 
mus was unfortunately used. Briefly: in 1928 the International (iommission on Zoological 
Nomenclature placed Limulus of 0. F. Miiller 1785 on the "Official List of Generic Names 
in Zoology" in the mistaken belief that this name was available and valid. However, 
Xiphosura was later found to have been used by Brunnich in 1771 for this animal, and so 
had priority. The discovery that the 1928 Opinion was made in error led some workers 
to consider it "not binding", and in the period around 1950 the authors of several works 
(including "Selected Invertebrate Types") used Xiphosura . In 1955, the International 
Commission, acting under its plenary powers (suspension of the rules) invalidated the 
priority of Xiphosura as a generic name of the American horseshoe crab. Opinion 320, 
including the letters expressing the views of specialists, makes instructive reading. 

Class Pycnogonida 

Despite the abundance of pycnogonids (about 50 genera and 500 species) in the seas 
of the world, the fauna of Woods Hole includes only three common species: Tanystylum 
orbiculare , Callipallene brevirostris , and Anoplodactylus lentus . This group has re- 
ceived little attention in recent years, although the first good systematic report on 
them in America was that on New England Pycnogonida by E. B. Wilson, and their embry- 
ology was described by T. H. Morgan in his doctoral thesis (1891). Most shore pycno- 
gonids feed upon hydroids and the young stages of many species encyst or form galls 
in hydroids. Since our information is still incomplete, collectors would do well to 
note the associated coelenterate in making collections. The following key will sep- 
arate the common local species, plus one common north of the Cape; for anything that 
will not key out, consult Hedgpeth (1948). 

(Figure references are to Plate 11) 

1. Chelifores present; palpi lacking (fig. 1) 2 

1. Chelifores absent; palpi present, of 4-7 joints (fig. 4); 

small species (TANYSTYLIDAE) Tanystylum orbiculare 

2. Ovigerous legs 10- jointed and present in both sexes; 

(fig. 3) (PALLENIDAE) Callipallene brevirostris 

2. Ovigerous legs less than 10- jointed, and lacking in 

females (fig. 6) PHOXICHILIDIIDAE 3 



3. Cephalic segment extended forward as a short neck, 

overhanging base of proboscis (fig. 5) Anoplodactyius lentus 

3. Cephalic segment not forming a neck (fig. 2) (common 

north of Cape) Phoxichilidium femora turn 


Anoplodactyius lentus Wilson, 1878. Breeds in August at Woods Hole. Dawson has described 
the colored blood corpuscles (Biol. Bull., 66_: 1934). Rare north of Cape. In Morgan's 
embryological work, called Phoxichilidium maxillare . 

Callipallene brevirostris (Johnson, 1837) . The smallest of the common pycnogonids at 
Woods Hole. Found among hydroids and on pilings. Referred to by Morgan as Pallene 
em pus a . 

Endeis spinosa (Montagu, 1808) . Not in key. Occasional at Woods Hole upon drifting Sar - 
gassum . 

Pvcnogonum littorale (Strom, 1762). Not in key. No record for Woods Hole, although with- 
in the reported range. 

Phoxichi lidium femora turn (Rathke, 1799) . Has been taken abundantly on Tubularia north 
of the Cape. 

Tanvstylum orbiculare Wilson, 1878. Common but small and easily overlooked; found on 
pilings and among ascidians and hydroids. 


Hedgpeth, J. W. , 1948. The Pycnogonida of the western North Atlantic and the Caribbean. 

Proc. U. S. Nat. Mus., 9J7_: 157-342. 
Morgan, T. H. , 1891. A contribution to the embryology and phylogeny of the pycnogonids. 

Studies from the Biol. Lab., Johns Hopkins Univ., 5_: 1-76, pi. I -VIII. 
Wilson, E. B., 1878. Synopsis of the Pycnogonida of New England, Trans. Conn. Acad. 

Arts & Sci. , 5: 1-26. 


Plate 11 


(1-6) Pycnogonids after Hedgpeth (scale bars = 1 mm); 
(7-19 barnacles after Zullo; all redrawn by Bruce 

Shearer . 

1. Anatomy of a generalized pycnogonid: Abdomen (ab) 

Eye tubercle (et) 

2. Phoxichilidium f e moratum . Chelifore (ch) 

Palp (pa) 

3. Callipallene brevirostris . Proboscis (pr) 

Ovigerous leg (ov) 

4. T anystylum orbicular e. Femur (f) 

Tarsus (t) 

5. Anoplodactylus lentus . Tibia, first, second (t-^, t 2 ) 

Propodus (p) 

6. Ovigerous leg of male A. lentus ■ 

7. Generalized lepadomorph barnacle with capitular sheath cut away to show 
cirri and filamentary appendages. 

8. Balanus balanus , base of shell wall seen from below, showing internal 
ribs and septa. 

9. Balanus improvisus , shell only, showing radius only partly overlapping 

10. Balanus eburneus , shell only, show- ala (a) 

ing extensive overlap of ala by carina (car) 
radius. carinolateral plate (clp) 

cirri (c) 

11. Lepas anserifera , seen from right excavation of tergum (e) 
side. filamentary appendages (f) 

lateral plate (lp) 

12. Diagram of plate arrangement paries (p) 
in Chthamalu s. radius (r) 

rostral plate (rp) 

13. Diagram of plate arrangement (rp = fused ros f rip) 
in Balanus . rostrolateral plate (rip) 

rostrum (ros) 

14. Balanus amphi trite niveus , showing scutum (s) 
color pattern of longitudinal striae. tergal spur (ts) 

tergum (t) 

15. Exterior of tergum of Balanus impro- 
visus , showing tergal spur. 

16. Exterior of scutum of _B. eburneus , showing radial striations. 

17. Interior of tergum of _B. eburneus , showing excavation on carinal side 
of basal margin. 

18. Interior of scutum of _B. amphi trite a mph i trite , showing adductor ridge 
diagonal line near center) . 


88 Chapter XI 


Class Crustacea 

Lower Crustacea and Cirripedia 

Keys are available only for some of the numerous and varied crustacean groups. 
Branchiopods in general are found in fresh water, and may be worked out with the 
aid of Ward and Whipple or of Pennak ' s guide to fresh water life. Copepods and 
ostracods, because of the number of species and their small size, do not lend them- 
selves to keying at the level of this guide. Cirripedes, amphipods , and isopods 
will prove difficult. The decapods, on the other hand, are well known and large 
forms, and, with some exceptions, can be identified with a fair degree of certain- 
ty. A key to the subclasses and major orders of crustaceans is not presented; the 
reader is referred to a basic reference text on invertebrate zoology. 

In matters of classification and nomenclature, these keys follow the system 
outlined by Waterman and Chace in Chapter I, Vol. I, of "The Physiology of Crus- 
tacea", edited by T. H. Waterman. The excellent systematic indices of the two 
volumes are very helpful in respect to the synonymy of experimentally used crusta- 
ceans . 

Subclass Cephalocarida (no key) Hutchinsoniella macracantha Sanders, 1955 is found 
in soft sediments at 10-30 meters depth. Not rare but very small (2-3.5 mm). 
See figure in Sanders, H. L., 1963. Cephalocarida. Functional morphology, lar- 
val development, comparative external anatomy. Mem. Conn. Acad. Arts & Sci., 
15 : 1-80. 

Subclass Branchiopoda 

Order Anostraca (no local marine representatives) 
Order Notostraca (no local marine representatives) 
Order Conchostraca (no local marine representatives) 

Order Cladocera (no key) : The genera Evadne and Podon occur in the marine plank- 
Subclass Ostracoda (no key) : Numerous marine representatives. 

Subclass Mystacocarida (no key) : Derocheilocaris tvpicus Pennak and Zinn (Smithson- 
ian Misc. Coll., 103(9): 1943) is the type of this subclass. It is minute (0.4- 
0.5 mm) and lives in the spaces between sand grains on Nobska and Falmouth bea- 

Subclass Copepoda (no key) : Extremely numerous in sea, both free living and para- 
sitic. See Wilson, C. B. , 1932. The copepods of the Woods Hole Region, Massa- 
chusetts. U. S. Nat. Mus. Bull. 158 : I-XIX, 1-635. 

Subclass Branchiura (no key): "Fish lice"; the genus Argulus occurs on fish in 

both fresh and salt water. Formerly considered an order (Arguloida) of the Co- 
pepoda, but now separated on the basis of their compound eyes and other fea- 
tures. For local species see Wilson (1932; cited above) pp. 11-18. 

Subclass Cirripedia 89 

by Victor A. Zullo 

Order Thoracica 
(Figure references are to Plate 11) 

Classification and identification of thoracic barnacles is based primarily on 
three sets of criteria: 1) mode of attachment to the substrate, 2) number, morpho- 
logy and arrangement of the calcareous plates forming the shell wall ( compartmental 
plates ) and closing the orifice ( opercular valves ) , and 3) morphology of the mouth 
parts ( trophi ) and appendages ( cirri ) . The Thoracica of this region are easily di- 
vided into two suborders on the basis of the mode of attachment. Members of the Sub- 
order Lepadomorpha are distinguished by the presence of a conspicuous, fleshy attach- 
ment stalk ( peduncle ) below the distal, expanded end ( capitulum ) which is more or less 
completely sheathed by calcareous plates. The shell in members of the Suborder Bal- 
anomorpha, on the other hand, is attached directly to the substrate, and it is to 
this group that most of the common barnacles of the Cape Cod Region belong. 

The lepadomorphs included in the key are all of the genus Lepas . Specific deter- 
mination is based 1) on the form and ornamentation of the plates ( scutum , terqum . and 
carina ) sheathing the capitulum (fig. 11), and 2) on the number of filamentary appen- 
dages present at the base of Cirrus I and on the prosoma. This latter feature can 
only be observed by removing one side of the capitular sheath to expose the body of 
the barnacle (fig. 7). 

The balanomorphs commonly found in this region ( Chthamalus and Balanus) have a 
shell wall composed of six compartmental plates and the orifice filled by four oper- 
cular valves (2 scuta and 2 terga) (figs. 12-13). Both end plates of the shell wall 
( rostrum and carina ) are overlapped by the adjacent compartmental plates in the genus 
Chthamalus (fig. 12), but in the genus Balanus , one of the end plates ( rostral plate ) 
overlaps the adjacent plates (fig. 13). 

Specific identification of balanomorph species requires a more detailed examina- 
tion of the opercular valves and shell. In some species, the basal plate sealing the 
barnacle to the substrate is membranous, but in others a calcareous plate is laid down 
and often remains attached to the substrate upon removal of the barnacle. The shell 
wall in many species of Balanus is formed of vertical inner and outer laminae separ- 
ated by longitudinal septa. These septa are usually reflected on the inner shell wall 
as regularly spaced, longitudinal ribs, which are especially evident near the base of 
the shell wall. In some instances the number of ribs exceeds the number of septa, and 
this character is used to differentiate certain species (fig. 8). The development of 
the articulation between adjacent compartmental plates is also a useful character in 
the identification of some taxa. In B. improvisus , for example, the overlapping sec- 
tion of the plate ( radius , plural radii ) is narrow and does not cover the overlapped 
section (ala, plural alae ) or extend to the central part of the adjacent plate ( par - 
ies , plural parieties ) (fig. 9). 

The removal and examination of opercular valves is necessary for proper identifi- 
cation of most species. The presence or absence on the scutum of an adductor ridge, or 
external radial striae (figs. 16, 18), and the degree of development of the tergal spur 
(figs. 15, 17) are characters which are used in the key. However, there are many oth- 
er features of these valves which are equally significant in specific determination. 


1. Animal enclosed in shell composed of varying numbers of plates 2 

1. Animal without shell, found in burrow with comma shaped aper- 
ture in the shells of gastropods occupied by hermit crabs . . Trypetesa lampas 

2. Shell attached directly to substrate Suborder BALANOMORPHA 3 

2. Shell attached to substrate by fleshy stalk or peduncle Suborder LEPADOMORPHA 9 

90 Cirripedia 

3. Both end plates of shell wall (rostrum and carina) over- 
lapped by adjacent plates (fig. 12); shell dull brown or 
grey; basis membranous Chthamalus fragilis 

3. Only one end plate (carina) overlapped by adjacent compart- 
mental plates, the opposing rostral plate overlapping ad- 
jacent plates (fig. 13) Balanus 4 

4. Basis membranous Balanus balanoides 

4. Basis calcareous 5 

5. Exterior of scutum distinctly striate longitudinally (fig. 16) 6 

5. Exterior of scutum lacking distinct longitudinal striae 7 

6. Ribs at base of interior of parieties more numerous than 
parietal septa (fig. 8); basis solid; basal margin of tergum 

entire Balanus balanus 

6. Ribs at base of interior of parieties a continuation of par- 
ietal septa; basis porose; carinal side of basal margin of 

tergum deeply excavated (fig. 17) Balanus eburneus 

7. Adductor ridge of scutum absent Balanus crenatus 

7. Adductor ridge of scutum well developed (fig. 18) 8 

8. Radii not extending to parieties of adjacent compartmental 
plates, with oblique summits (fig. 9); tergal spur narrow, 
less than V4 width of basal margin (fig. 15); parieties white 

Balanus improvisus 

8. Radii extending to parieties of adjacent compartmental plates, 
with slightly sloping summits (fig. 10); tergal spur at least 
l/l width of basal margin; exterior of parieties with regular- 
ly arranged gray, purple, or red-purple longitudinal striae 

(fig. 14) Balanus amphi trite niveus 

9. Carina terminating below in an expanded disk, with umbo pro- 
jecting angularly; valves thin, papery Lepas fascicularis 

9. Carina terminating below in a fork; umbo basal; valves well 

calcified 10 

10. Valves radially furrowed or strongly striate (fig. 11) 11 

10. Valves smooth or minutely striate 12 

11. Occludent margin of scutum arched, protuberant, forming wide 
shelf between margin and umbonal-apical ridge; 5-6 filamentary 

appendages on either side of body Lepas anserifera 

11. Occludent margin of scutum not arched, nearly parallel with 
umbonal-apical ridge, leaving comparatively narrow area be- 
tween margin and ridge; 0-2 filamentary appendages on either 

side of body at base of Cirrus I Lepas pectinata 

12. One or 2 filamentary appendages (fig. 7) on either side of 

body Lepas anatifera 

12. Three filamentary appendages on either side of body 

Lepas hilli 

Cirripedia 91 


Order Thoracica 

Suborder Lepadomorpha 

Lepas anatif era Linnaeus, 1758. Common during July and August on floating wood and Sar - 
gassum . 

Lepas anserif era Linnaeus, 1767. Occasionally associated with L. pectinata on drift- 

Lepas fascicularis Ellis and Solander, 1786. Found attached to floating Sargassum dur- 
ing the summer months. 

Lepas hilli Leach, 1818. Some of the records of this species from the Cape Cod Region 
may be based on misidentif ications of L. anatif era . 

Lepas pectinata Spengler, 1793. This species is often associated with L_. anatifera on 
floating wood in Buzzards Bay and Vineyard Sound during the summer months. 

Suborder Balanomorpha 

Balanus ( Balanus ) amphi trite niveus Darwin, 1854. ( = B. venustus niveus ) . The most 

common barnacle in the subtidal waters of Buzzards Bay and Vineyard Sound. Found 

on stones and shells, especially those of gastropods occupied by hermit crabs. Not 

known north of Cape Cod nor in Cape Cod Bay. 
Balanus ( Balanus ) balanus (Linnaeus 1758) . (- B. porcatus ) . A large, strongly ribbed 

barnacle in the lower intertidal zone north of and throughout the Cape Cod Canal. 

Occasional specimens are found subtidally in Vineyard Sound. 
Balanus (Balanus ) crenatus Brugui&re, 1789. Occasionally associated with B. amphi - 

trite niveus in the subtidal waters of Vineyard Sound, but more common in Cape Cod 

Balanus ( Balanus ) eburneus Gould, 1841. Common in the lower intertidal of protected 

inlets and bays, and especially in waters of lower salinity with B. improvisus . 
Balanus (Balanus ) improvisus Darwin, 1854. Common in waters of low salinity on the south- 
ern coast of Cape Cod, and often associated with B. eburneus . 
Balanus ( Semibalanus ) balanoides (Linnaeus, 1767). The most abundant and ubiquitous 

barnacle in the intertidal zone of the Cape Cod Region, but drops out in brackish 

waters . 
Balanus ( Chirona ) hameri (Ascanius , 1761) . Not in key. Common on the large commercial 

scallop, Placopecten magellanicus , abundant in deeper water, as on fishing banks. 

Potentially important to physiologists because it is by far the largest barnacle 

in this region, over 5 cm in basal diameter. 
Chthamalus f ragilis Darwin, 1854. Abundant in the upper intertidal zone of Buzzards 

Bay and Vineyard Sound. Also present in limited numbers along the southern shore 

of Cape Cod Bay, but not known to occur north of this area. 

Order Acrothoracica 

Trypetesa lampas (Hancock, 1849) . (= Alcippe lampas ) . Found only in the floor and side 
of the interior of the body whorl of gastropod shells such as Lunatia heros and Ne - 
verita duplicata occupied by hermit crabs. 


A monograph on the sub-class Cirripedia, Lepadidae. Ray Society, 
1-400, pis. 1-10. 
A monograph on the sub-class Cirripedia, Balanidae, Verrucidae. 
Ray Society, London, i-viii, 1-684, text figs. 1-11, pis. 1-30. 
Genthe, K. W. , 1905. Some notes nn Alrippp lampas. and its occurence on the American 
Atlantic shore. Zool. Jahrb. (Anat.), 21: 181-200. 

Darwin , C . , 




Darwin, C . , 


92 Cirripedia 

Pilsbry, H. A., 1907. The barnacles (Cirripedia) contained in the collections of the 
U. S. National Museum, Bull. U. S. Nat. Mus. 60: i-x, 1-122, text figs. 1-36, pis. 

Pilsbry, H. A., 1916. The sessile barnacles (Cirripedia) contained in the collections 
of the U. S. National Museum; uncluding a monograph of the American species. Ibid . , 
93 ; i-xi, 1-366, text figs. 1-99, pis. 1-76. 

Zullo, V. A., 1963. A preliminary report on systematics and distribution of barnacles 
(Cirripedia) of the Cape Cod region. Systematics-Ecology Program, Marine Biologi- 
cal Laboratory, Woods Hole, Mass., 33 pp. 

Order Rhizocephala 

The Rhizocephala are parasitic cirripeds which infest decapod crustaceans and iso- 
pods. The adult rhizocephalan is highly modified in body form and bears no resemb- 
lance to other cirripeds. The affinities of this group are exhibited only by the lar- 
val stages which include a nauplius with characteristic frontal horns (although lack- 
ing an alimentary canal) and a cypris. 

The life history of the common European species Sacculina carcini (Thompson) was 
worked out by G. Smith in 1907. The fertilized eggs mature and develop into cirri- 
pedan nauplii within the "mantle cavity" of the parent. The nauplii are released and 
swim actively for four days during which they pass through four molts. On the fifth 
day the nauplius undergoes metamorphosis into the cypris stage which is also active- 
ly free-swimming and lasts from two to three days. About the third day the cypris 
attaches itself by the antennules to the base of a seta of the portunid crab Car - 
cinus maenas . The thoracic appendages and musculature are cast off and a new larva, 
the kentrogen, composed of a mass of mesodermal cells surrounded by an ectodermal 
chitinous bag is produced under the old cypris shell. A hollow, dart-like ectoderm- 
al process pierces the base of the seta to which the kentrogen is attached, and the 
enclosed mesodermal cells pass through into the haemocoel of the crab and settle near 
the midgut. The cells rapidly divide and form a tumor about the midgut with numer- 
ous divergent roots which eventually extend into the extremities of the crab. The 
main body mass of Sacculina emerges from the interior after the following molt of 
the host at a point near the ventral juncture of the thorax and abdomen. Further 
molting is inhibited by the parasite, whose adult organs (including paired ovaries 
and testes, and a neural ganglion) now differentiate. 

Rhizocephalans have not been reported from the Cape Cod Region, although Sac - 
culina carcini parasitizes the green crab Carcinus maenas , and Peltogaster the her- 
mit crab Pagurus bernhardtus in European waters. The works of Erof essor Hilbrand 
Boschma (1925-1963) may be consulted for details on the Rhizocephala. A complete 
bibliography is given in Zool. Meded. (1964), vol. 39, pp. XLI-XLVI. 

Chapter XII 93 


Class Crustacea 

The Lower Malacostraca (Peracarida) 

The successful, abundant, and varied peracaridan crustaceans are most commonly 
represented in shore collecting by isopods and amphipods, the general features of 
which are well known to the average zoologist. In their rather specialized morpho- 
logy these orders depart somewhat from the more generalized and shrimp-like body 
form which is basic to the Subclass Malacostraca. However, certain other peracari- 
dan orders, the swimming Mysidacea and the small, bottom-dwelling Cumacea, retain 
an essentially shrimp-like form, with carapace and elongated abdomen. The Key to 
Orders given below will aid in the separation of these groups. The diverse peracari- 
dan orders share one feature, namely, the brood pouch or "marsupium" formed of plates 
(oSstegites) borne by the thoracic legs of the female, and used to house the eggs and 
developing young. 


1. Body having the "caridoid" (shrimplike) form, with a dis- 
tinct carapace over the thorax and an elongated abdomen 2 

1. Body having thorax and abdomen not sharply distinguishable; 

carapace lacking or very small 3 

2. Eyes stalked when present; carapace covering all or most of 

thorax; swimming forms MYSIDACEA (p. 93 ) 

2. Eyes sessile when present; carapace covering only 3 or 4 
thoracic segments and inflated into a branchial chamber on 

each side; in bottom sediments CUMACEA (p. 98 ) 

3. A small carapace present, covering only 2 thoracic segments; 
resemble small isopods but have first pair of legs chelate 


3. Carapace lacking 4 

4. Body usually dorsoventrally flattened; thoracic legs (except 
for maxilliped) essentially alike; abdomen with 5 pairs of 

pleopods with unsegmented rami, and 1 pair of uropods ISOPODA (p. 102) 

4. Body usually laterally compressed; thoracic limbs of more 
than one form, with the 2nd and 3rd pairs usually prehen- 
sile; abdominal appendages consist of 3 pairs of pleopods and 
3 pairs of uropods AMPHIPODA (p. 107) 


By Roland L. Wigley 

The mysids are the most typically shrimp-like ('caridoid") of the various pera- 
caridan orders. They are adapted for swimming, with elongated bodies and a well dev- 
eloped carapace covering almost the entire thorax. The lightly calcified integument 
is thin and flexible. The antennules are biramous, having multi-segmented flagellar 
antennae have an exopod, usually in the form of a flattened scale (antennal scale) ; 
the endopod is f lagelliform, composed of numerous segments. Eyestalks are cylindri- 
cal, well developed, terminating in a prominent brown or black (rarely red) cornea. 
The eight thoracic segments each bear biramous appendages; branchiae are absent in 
the species listed here. Females bear a large marsupium (hence the common name 
Opossum Shrimp) composed of 2-7 pairs of oostegites attached to the thoracic limbs. 
The pleopod structure varies, depending on sex and species. 

94 Mysids 

Mysids inhabit a wide variety of benthic and planktonic habitats. They are com- 
mon in brackish and marine waters, but no freshwater forms are known in this region. 
Mysids are an important link in littoral and continental shelf food webs, and are par- 
ticularly valuable as food for small species of fish as well as the young of larger 
fishes. Seasonal inshore-offshore migrations and other horizontal movements have been 
observed for a few species. Pronounced vertical migrations in which they swim upward 
during hours of darkness are characteristic of this group. 

(Figure references are to Plate 12) 

1. Telson with posterior end cleft (figs. 4, 8, 9, 11) 2 

1. Telson with posterior end entire (figs. 5-7, 10) 5 

2. Antennal scale with setae present on both inner and outer 

margins (fig. 2) 3 

2. Antennal scale with outer margin devoid of setae and termin- 
ating in an articulated spine (fig. 4) Praunus f lexuosus 

3. Telson with the entire lateral margins armed with spines. 
Antennal scale elongate, width 9 to 12 times length, produced 

into an acute apex (figs. 8, 9) 4 

3. Telson with basal one-half of the lateral margins devoid of 
spines; antennal scale elliptical, apex blunt, 3.5 times as 

long as broad (figs. 3, 11) Heteromysis formosa 

4. Antennal scale approximately 9 times as long as broad, outer 
margin nearly straight, telson lateral margin spines more than 

30, extending posteriorly to apex (fig. 8) Mysis mixta 

4. Antennal scale approximately 12 times as long as broad, outer 
margin concave in outline; telson lateral margin spines 25, ex- 
tending posteriorly as far as the cleft (fig. 9) Mysis stenolepis 

5. Antennal scale with setae present on both inner and outer mar- 
gins; telson lateral margins armed with numerous spines (figs. 

7, 10) 6 

5. Antennal scale with setae absent on outer margin, terminating in 
a strong spine; telson without spines on lateral margins (figs. 

5, 6) 7 

6. Antennal scale about 10 times as long as broad (fig. 10); telson 

subequal in length to endopod of the uropod Neomysis americana 

6. Antennal scale about 5 times as long as broad (fig. 7); endopod 

of the uropod 1.5 times as long as the telson Mysidopsis bigelowi 

7. Eyes dorso-ventrally flattened, cornea kidney-shaped, red color; 
telson length subequal to its greatest width (fig. 5) 

Erythrops ery throphtha lm a 

7. Eyes globular not dorso-ventrally flattened, cornea black; 
telson elongate, length 2 times its greatest width (fig. 6) 
Meterythrops r obusta 


Erythrops ery throphtha lma (Goes, 1863). Occasionally referred to as Erythrops 

goesii . Inhabits deeper waters, 40-2 75 meters, from Cape Cod to the Arctic. One 
of the most beautifully colored New England mysids. Eyes brilliant carmine red; 
opaque white pigment spots scattered over the body; an orange-red dorsal patch 

Mysids 95 

on the carapace and clear bright yellow pigment spots on the ventral body surface. 
Adult length 9-11 mm. 

Heteromysis formosa S. I. Smith, 1873. A common species from New Jersey to Canada. 

Frequently in small swarms inside dead bivalve shells such as Spisula . Intertid- 

al to 248 meters. Males semi-translucent, but parts of the females are a rose 
color. Adult length 6-9 mm. 

Metervthrops robusta S. I. Smith, 1879. Uncommon; in the western Atlantic, occurs 
from Cape Cod to Greenland. 66-300 meters. Adult length 14-16 mm. 

Mysidopsis biqelowi Tattersall, 1926. A warm water species, from Louisiana to Cape 
Cod. Most common at shallow shelf depths, 16-50 meters, but has been found to 
196 meters. Adult lenqt-h 7.5 mm or somewhat more. 

Mysis mixta Lilljeborg, 1852. Occasionally referred to as Michtheimysis mixta . 

A common species of the east coast from Woods Hole to Canada. Closely related to 
Mysis stenolepis ; however, in contrast, M. mixta inhabits areas where algae and 
Zostera are absent. Intertidal to 200 meters. Adult length 20-25 mm. 

Mysis stenolepis S. I. Smith, 1873. Sometimes referred to as Michtheimysis stenolep - 
is and Mysis spinulosus . Inhabits intertidal and shallow shelf waters from New 
Jersey to Gulf of St. Lawrence. Closely related to M. mixta (see above), but is 
a more strictly littoral weed inhabitinq species. Adult length 25-30 mm. 

Neomysis americana (S. I. Smith, 1873). Mysis americana is a synonym. Very common, 
from Virginia to the Gulf of St. Lawrence. Intertidal to 214 meters. Adult 
length 10-12 mm. 

Praunus f lexuosus (Muller, 1776) . A European species of relatively large size, first 
found in American waters in 1960. Has been reported only in the harbor at Barn- 
stable, Mass. In Europe is one of the commonest species in brackish waters and 
tidal zones along the coast. Adult length 24-25 mm. 


Lochhead, J. H., 1950. Heteromysis formosa . In : Selected Invertebrate Types. Frank 
A. Brown, Jr., ed. John Wiley and Sons, Inc., N. Y. 

Smith, S. I., 1879. The stalk-eyed crustaceans of the Atlantic coast of North America 
north of Cape Cod. Trans. Conn. Acad. Arts and Sci., 5_: 27-138, pis. 7-12. 

Tattersall, W. M. , 1951. A review of the Mysidacea of the United States National Mu- 
seum. U. S. Nat. Mus. Bull. 201 . : I-X, 1-292. 

Tattersall, W. M. , and Olive S. Tattersall, 1951. The British Mysidacea. Ray Society, 

Verrill, A. E. , S. I. Smith, and O. Harger, 1873. Catalogue of the marine inverte- 
brate animals of the southern coast of New England, and adjacent waters. Sec- 
tion D. Ijn: Report upon the invertebrate animals of Vineyard Sound and adjacent 
waters, with an account of the physical characters of the region. By A. E. Ver- 
rill. Rept. U. S. Comm. Fish and Fisheries, 1871-72: 537-749, pis. 1-38. 


Plate 12 


Figures mainly from Tattersall (1951) and 
Tattersall and Tattersall (1951), redrawn 
by Ruth L. von Arx. 

Fig. 1. Side view of a typical mysid, ovigerous female, thoracic 
legs 1-7 omitted. 

2. Antennal scale and basal portion of antenna of a typical mysid. 

3. Heteromysis formosa , dorsal view. 

4-11. Upper row shows telsons, lower row shows antennal scales, 
both in dorsal view, of the following species: 

4 . Praunus f lexuosus . 

5 . Erythrops erythrophathalma . 

6 . Meterythrops robusta . 

7. Mysidopsis bigelowi . 

8. Mysis mixta . 

9. Mysis stenolepis . 

10. Neomysis americana . 

11. Heteromysis formosa . 

Plate 12 

cornea eyestalh 

horacic somites ( I - 8) 

abdominal somites (l — 6) 




10 II 


f leiuosus 















98 Cumaceans 


By Roland L. Wig ley 

Cumaceans, although widespread in marine situations and with some species in 
brackish waters, are not well known to the average zoologist because of their small 
size and habit of living buried in sand or mud. The general body form is distinc- 
tive, characterized by an inflated cephalothorax and a slender, cylindrical abdomen. 
The carapace is moderately large, covering the anterior half of the cephalothorax. 
The integument in many species is rather heavily calcified and brittle. The first 
thoracic limb (maxilliped) bears a membranous epipodite, usually furnished with res- 
piratory lamellae. A pseudorostrum is present in many species, formed by carapace 
lateral plates coming together above and in front of the head. Eyes are present in 
some species, sessile and usually coalesced to form a single organ on a lobe located 
mid-dorsally on anterior portion of carapace. First antenna with inner flagellum 
usually reduced or absent; second antenna vestigial in females, well developed in 
males. First 3 thoracic appendages modified as maxillipeds; remaining 5 as peraeo- 
pods. One pair of large styliform uropods present. Telson is distinct, or coa- 
lesced with last abdominal somite. Adults in this region generally range from 3 
to 15 mm in length. 

There is pronounced sexual dimorphism; pleopods are absent in all females and 
in males of some species, but males generally have from 1 to 5 pairs. Eggs are in- 
cubated in a marsupium, and hatch as postlarvae in which the past pair of peraeo- 
pods is undeveloped. 

Cumaceans are primarilv marine, but a few species occur in brackish waters. They 
are widely distributed in all oceans from intertidal zone to great depths in mid- 
ocean, most common subtidally. Basically benthic in habit, they commonly burrow in 
mud or sand. They occasionally undertake pelagic excursions, chieflv at night, at 
which time males greatly outnumber females. 

(Figure references are to Plate 13) 

1. Independent telson present (figs. 23-26) 2 

1. Without independent telson (figs. 15-22) 7 

2. Uropod inner ramus 1- jointed; telson without spines 

(figs. 13, 23) Petalosarsia declivi s 

2. Uropod inner ramus 2- or 3- jointed; telson with lateral and 

apical spines (figs. 24-26) 3 

3. Telson with 5 apical spines; male without pleopods . . . Lamprops quadriplicata 

3. Telson with or 2 apical spines; male with 2 pairs of pleo- 
pods Family Diastvlidae 4 

4. Telson apex upturned, without apical spines (figs. 11, 26) 

Oxyurostylis smithi 

4. Telson apex not upturned, 2 apical spines (figs. 24, 25) Diastylis 5 

5. Carapace with 4 large spines on dorsal surface (figs. 1, 24) 

Diastylis quadrispinosa 

5. Carapace without dorsal spines (figs. 6, 7) 6 

6. Carapace with 3 oblique lateral ridges; horizontal ridge on 
postero-lateral section of carapace in male (fig. 6) Diastylis p olita 

6. Carapace with 4 oblique lateral ridges; without horizontal 
ridge on postero-lateral section of carapace in male (figs. 
7 25) Diastylis sculpta 

Cumaceans 99 

7. Mandibles broadly truncate at base (fig. 14) ; male with 2 

pairs of pleopods Family Leuconidae 8 

7. Mandibles narrow or acuminate at base (fig. 13); male with 

0, 3, or 5 pairs of pleopods 12 

8. Pseudorostrum well developed Leucon americanus 

8. Carapace truncate anteriorly without pseudorostrum 9 

9. Uropods with outer ramus longer than inner ramus (fig. 18) 

Eudorellopsis deformis 

9. Uropods with outer ramus shorter than inner ramus (figs. 

15, 16) Eudorella 10 

10. Body setose; approximately 14-18 teeth on carapace ventral 

margin (figs. 8, 15) Eudorella hispida 

10. Body not setose; less than 14 teeth on carapace ventral margin 11 

11. Anterior margin of carapace deeply emarginate in female, 

slightly emarginate in male (figs. 2, 3, 20) Eudorella emarqinata 

11. Anterior margin of carapace slightly emarginate in female, 

serrate or entire in male (figs. 9, 16) Eudorella truncatula 

12. Endopod of uropod 2- jointed; 3 pairs of pleopods in male 

(figs. 10, 17) Leptocuma minor 

12. Endopod of uropod 1- jointed; or 5 pairs of pleopods in 

male (figs. 19, 22) 13 

13. Male has no pleopods and a rudimentary second antenna; in 
female, uropod peduncle length about equal to maximum width 

of last abdominal somite (figs. 4, 22) Almyracuma proximoculi 

13. Male has 5 pleopods and well developed second antenna; in fe- 
male, uropod peduncle length greater than 2 times maximum width 
of last abdominal somite (figs. 5, 19) Cyclaspis varians 


Almyracuma proximoculi Jones, 1959. Brackish water species, known only from Pocas- 

set River. Adult size 3-4 mm. 
* Campy las pis af finis G. 0. Sars, 1869. Not yet reported from Cape Cod waters, but 

expected to occur in deeper portions of this area. 
* Campylaspis rubicunda (Lilljeborg, 1855) . Not yet reported from Cape Cod waters, 

but expected to occur in the deeper portions of this area. 
Cyclaspis varians Caiman, 1912. Brackish water species found in surface waters of 

Woods Hole Harbor and Vineyard Sound. Adult size 3-4 mm. 
* Diastyli s abbreviata G. 0. Sars, 1871. Not yet reported from Cape Cod waters, but 

expected to occur in deeper portions of this area. 
Diastylis polita S. I. Smith, 1879. Found at depths less than ca. 35 meters from 

Nova Scotia to Block Island Sound. Adult size 12-14 mm. 
Diastylis quadrispinosa G. 0. Sars, 1871. Occurs from the Gulf of St. Lawrence to 

New Jersey in depths of 4-400 meters. Adult size 11 mm. 
Diastylis sculpta G. 0. Sars, 1871. In American waters occurs from Gulf of St. 

Lawrence to Long Island; low water to ca. 400 meters. Adult size 9-10 mm. 
Eudorella e marginata (Kroyer, 1846). In American waters occurs from the Arctic to 

Martha's Vineyard in depths from 2-130 meters. Adult size 10-12 mm. 
Eudorel la h ispida G. 0. Sars, 1871. Occurs from the Arctic to Martha's Vineyard, 

in depths from 30-130 meters. Adult size 5-6 mm. 
Eudorella truncatula (Bate, 1856). On U. S. coast reported from Massachusetts Bay 

to Block Island Sound, in depths of 30-100 meters. Adult size 4-5 mm. 


Plate 13 


Figures mainly from Sars (1900) and Caiman 
(1912), redrawn by Ruth L. von Arx. 

Fig. 1. Diastylis quadrispinosa , female, dorsal view. 

2. Eudorella emarqinata, female, lateral view. 

3. Eudorella emarqinata , male, lateral view 

4-12. Carapace in lateral view of the following species: 

4. Almyracuma proximoculi , male. 

5. Cyclaspis varians , female. 

6. Diastylis polita , male. 

7. D iastylis sculpta , male. 

8. Eudorella hispida , female. 

14. Eudorellopsis deformis , mandible. 

15-26. Dorsal view of last abdominal somite, uropod, and telson of the 
following species: 

9. Eudorella truncatula , female. 

10. Leptocuma minor , male. 

11. Oxyurostylis smithi , male. 

12. Petalosarsia declivis, male. 

13. Petalosarsia declivis , mandible. 

15. Eudorella hispida , female. 

16. Eudorella truncatula , female. 

17. Leptocuma minor , female. 

18. Eudorellopsis deformis , female. 

19. Cyclaspi s varians , male. 

20. Eudorella emarginata, female. 

21. Leucon americanus , male. 

22. Almyracuma proximoculi , male. 

23. Petalosarsia declivis , female 

24. Diastylis quadrispinosa , female. 

25. Diastylis sculpta , female. 

26. Oxyurostylis smithi, female; 
(inset: telson tip in side view) 

Plate 13 

102 Cumaceans 

Eudorellopsis deformis (Kroyer, 1846) . In American waters reported from the Arctic 
to Long Island in depths of ca. 2-100 meters. Adult size 4-5 mm. 

Lamprops quadriplicata S. I. Smith, 1879. Occurs from Newfoundland to southern New 
England, in depths from ca. 2-150 meters. Adult size 8-9 mm. 

Leptocuma minor Caiman, 1912. Occurs from Gloucester, Mass. to Woods Hole, in depths 
of ca. 2-100 meters. Adult size 6-7 mm. 

Leucon americanus Zimmer, 1943. A brackish water species reported from Woods Hole 
and Chesapeake Bay. Adult size 4-5 mm. 

Oxyurostylis smithi Caiman, 1912. A euryhaline species, Maine to Louisiana, from 
surface to 20 meters. Adult size 6-7 mm. 

Petalosarsia declivis (G. 0. Sars, 1864) . In American waters occurs from the Arc- 
tic to Buzzards Bay, in 20-200 meters. 

*Not in key. 


Caiman, W. T., 1912. The Crustacea of the order Cumacea in the collection of the 

United States National Museum. Proc. U. S. Nat. Mus . 41: 603-676, text figs. 

Jones, N. S. and W. D. Burbanck, 1959. Almyracuma proximoculi gen. et sp. nov. 

(Crustacea, Cumacea) from brackish water of Cape Cod, Massachusetts. Biol. 

Bull. 116: 115-124.. 
Sars, G. 0., 1900. An account of the Crustacea of Norway, vol. 3, Cumacea, pp. 

I-X, 1-115, pis. 1-72. 
Smith, S. I., 1879. The stalk-eyed crustaceans of the Atlantic coast of North 

America north of Cape Cod. Trans. Conn. Acad. Arts and Sci. 5_: 28-138, pis. 



Isopods are readily recognizable in most instances, except for the grossly modi- 
fied parasitic forms. Since isopods are so common they are of great ecological sig- 
nificance, and certain species have been widely used in experimental work, hence 
their identification is a practical matter of much importance. Fortunately, the 
group is reasonably well known in the Woods Hole region, and specific identifica- 
tion does not offer such difficulties to the average zoologist as does the identi- 
fication of amphipods . Tanaidaceans, because of their small size, are less fre- 
quently noticed. Since they have often been classified as a group within the Iso- 
poda, and since they are so similar in general form, the two orders are combined 
in the following key. The advice of Thomas E. Bowman and of Roland L. Wigley in 
the preparation of the following key and check-list is gratefully acknowledged. 
Figure references on isopods and tanaidaceans are to Plates 14 and 15. 


First pair of legs chelate (figs. 1-3); 6 free thoracic seg- 
ments; tiny creatures Order TANAIDACEA (CHELIFERA) 

Not well known locally; possibly species include: 

a) 3 pairs of pleopods Tanais cavolinii 

b) 5 pairs of pleopods Leptochelia saviqnyi 

First pair of legs simple or subchelate; 7 free thoracic seg- 
ments Order ISOPODA 

Isopods 103 

2. Parasitic on Crustacea; body may be grossly modified; males 

tiny Suborder Epicaridea (Bopyroidea) 26 

2. Not parasitic on crustacea (a few are ectocommensal) ; body 

easily recognizable as that of an isopod 3 

3. Uropods terminal, set at rear (figs. 17-19, 29, 30) 22 

3. Uropods set laterally or ventrally (figs. 4-16, 26-28) 4 

4. Uropods lateral, visible from above, flattened, forming with 

telson a caudal fan in most instances 5 

4. Uropods ventral, not visible dorsally, turned inwards to form 

the cover of a chamber encasing the pleopods (fig. 11) . . .Suborder Valvifera 14 

5. Thorax with only 5 pairs of normal walking legs; not reported 

locally Suborder Gnathiidea 

5. Thorax with 7 pairs of more or less normal walking legs 6 

6. Exopodal part of uropod arched over telson; body very long 

(7 times the width) : Suborder Anthuridea; one species common 

locally in brackish situations (fig. 4) Cyathura polita 

6. Uropodal exopods lateral; body length seldom if ever more than 

5 times width Suborder Flabellifera 

7. Pleotelson (abdomen and telson) shows 6 segments (figs. 6-9) 10 

7. Pleotelson with segments fused so as to have only two complete 

segments, the terminal one of which is large and conspicuous 8 

8. Uropods each with two flattened branches; all legs simple; 

animals able to roll themselves into a complete ball 9 

8. Uropods each a single slender pointed process, lacking 
outer branches (exopods) ; terminal segment of abdomen tri- 
angular, with truncated apex (fig. 26); first and second 

pairs of legs subchelate Ancinus depressus 

9. Terminal segment of abdomen rounded at rear; exopods of uro- 
pods with outer serrated margin (fig. 27) Sphaeroma quadridentatum 

9. Terminal segment bluntly triangular; exopods of uropods with 

smooth outer margin (fig. 28) Exosphaeroma papillae 

10. Exopod of uropod flattened (not clawlike) and comparable 

to endopod in size, forming a normal caudal fan 12 

10. Exopod of uropod clawlike and much smaller than endopod 
(fig. 7); uropods very small relative to telson; animals 

burrow in wood ("gribbles") Limnoria 11 

11. Telson lacking tubercles and with a Y-shaped ridge (fig. 7) 

Limnoria liqnorum 

11. Telson with tubercles on posterior margin and with three tuber- 
cles on dorsal surface (fig. 8) Limnoria tripunctata 

12 . Bases of antennae seem to meet in front of the head as seen 
from above (fig. 5) ; head terminating in a small point; front 
3 legs with slightly hooked tips; endopod of uropod notched 

laterally Cirolana c oncharum 

12. Bases of antenna not meeting in midline (figs. 6, 9); all 
thoracic legs end in hooked dactyls; animals usually found 
clinging to fish or around fish-landing docks 13 

104 Isopods 

13. Form regular and oval; 2 longitudinal light stripes run 

length of body; head subquadrate with trilobed rear margin 

and distinct eyes; abdomen and uropods well developed; body 

to 13 mm long (fig. 9) Nerocila munda 

13. Form oval but with a slight asymmetry; head relatively small 
with eyes indistinct; abdomen with relatively small uropods; 

body stout, to 25 mm long (fig. 6) Lironeca ovalis 

14. Sides of head entire (figs. 10, 12); eyes lateral 18 

14. Sides of head notched or incised (figs. 23-25); eyes dorsal; 

body broad, with pointed telson Chiridotea 15 

(Note: see Bowman (1955) and Wigley (1961) for additional details 
on identification of Chiridotea species) . 

15. Second (more ventrally placed) antennae decidedly longer than 

the first pair (figs. 23, 24) 16 

15. Second antennae nearly equaling in length the first pair (fig. 

25) 17 

16. Sides of telson curved; finger of first large gnathopod with 
a single spine back of the terminal claw (fig. 23 A-C) ; ani- 
mal occurs in brackish water Chiridotea almyra 

16. Sides of telson straight, telson wedge shaped; finger of first 
gnathopod with several spines along the inner margin (fig. 24 

A-C) ; animal marine Chiridotea tuf tsi 

17. Clefts in sides of head deep (fig. 25A) ; antero-lateral margins 

of head with numerous bristles; body color light Chiridotea coeca 

17. Clefts in sides of head shallow; antero-lateral margins of head 
only sparsely provided with bristles; body color dark; occurs in 

brackish waters Chiridotea nigrescens 

18. Pleotelson seems to consist of 3 parts (plus grooves showing a 

4th partly coalesced segment) ; common Idotea 19 

18. Pleotelson seems to consist of one part (plus grooves suggest- 
ing one partly coalesced segment) 20 

19. Rear of telson bracket-shaped, with a central point (fig. 10) 
Idotea baltica 

19. Rear of telson truncated (cut square across) (fig. 12) . . Idotea me tallica 

19. Rear of telson pointed (fig. 13) Idotea phosphorea 

20. Pleotelson sides tapering to a point; in side view shows 2 
rounded elevations separated by a depression (fig. 14); sec- 
ond antennae short, reaching back only to second thoracic 

segment Edotea tri loba 

20. Pleotelson with more or less parallel sides cut off to a tri- 
angular point at rear; second antennae long, reaching to 5th 

thoracic segment Erichsonella 21 

21. Pleotelson with a pronounced tubercle on either side (fig. 15) 
Erichsonella f ilif ormis 

21. Pleotelson with only a suggestion of a tubercle on either side 

(fig. 16) Erichsonella attenuata 

Isopods 105 

22. Aquatic forms; pleotelson consisting of 2 sections .... Suborder Asellota 25 

22. Terrestrial or high-beach forms; pleotelson showing 

6 sections Suborder Oniscoidea 2 3 

(Most terrestrial forms are not in this key; members of the 
fully terrestrial genera Porcellio , Armadillidium , etc., may 
often be washed into bays and estuaries, to confuse the col- 
lector) . 

23. Less than 1 cm long; uropods decidedly shorter than pleotel- 
son 24 

23. Length may exceed 2 cm; uropods as long as pleotelson, each 
with 2 slender terminal articles (fig. 29) ; animals may be 
very active, running about like young cockroaches near high 

tide marks Liqia oceanica 

24. Under boards, rubbish, etc. above high tide marks; small 

(up to 6.5 mm long) ; body surface smooth (fig. 19) .... Philoscia vittata 

24. In beach sand; very small (up to 4.5 mm long); body sur- 
face thickly covered with small, spine-tipped tubercles 

(fig. 30) Scyphacella arenicola 

25. Uropods very small (fig. 18), in a posterior notch of 
telson; all thoracic legs simple; very small (to 5 mm long); 

in marine situations ffaera marina 

25. Uropods at least as long as telson (fig. 17); first legs sub- 
chelate; up to 15 mm long; in fresh water or water of quite 

low salinity Asellus sp. 

26. On Callianassa ; female with six pairs of jointed branching 
abdominal processes; small male with elongated appendages 

at sides of abdomen (fig. 20) lone thompsoni 

26. On Palaemonetes , forming a bulge in wall of gill chamber; 
female and male with short abdominal processes (fig. 21) 
Probopyrus pandalicola 

26. On Paqurus longicarpus ; female with pleopods and with 

6th thoracic segment longer than others; male with unseg- 

mented abdomen (fig. 22) Steqophryxus hyptius 

Order Tanaidacea (Chelifera) 

Leptochelia savignyi (Kroyer, 1842). 
Tanais cavolinii Milne-Edwards, 1829. 

Order Isopoda 

Suborder Anthuridea 

Calathura branchiata (Stimpson, 1853). Rare south of Cape. Not in key. 

Cyathura polita (Stimpson, 1855) . Common in areas of low or very low salinity, as 

in the tidal drainages from ponds. Until recently has been called C. carinata 

Kroyer (see Miller and Burbanck, 1961) . 
Ptilanthura tenuis Harger, 1879. Rare. Not in key. 

106 Isopods 

Suborder Flabellifera 

Ancinus depressus (Say, 1818) . Reported from Woods Hole (see Richardson, 1909) . 

Exosphaeroma papillae Bayliff , 1938. A brackish water species, reported from Long 
Island, and from Sandwich, on Cape Cod. 

Cirolana concharum (Stimpson, 1853). A scavenger, often very common around fish 
landing docks. 

Cirolana polita (Stimpson, 1853). Not in key. Reported north of Cape. 

Limnoria lignorum (Rathke, 1799) . Found burrowing into surface layers of sub- 
merged, untreated wood. 

Limnoria tripunctata Menzies, 1951. 

Lironeca ovalis (Say, 1818) . An ectoparasite on fish; uncommon. 

Livoneca , see Lironeca . The commonly used name Livoneca is the result of a typo- 
graphical error; the group of names including Lironeca , Cirolana , and Nerocila 
was intended as a series of anagrams on Caroline and Carolina. 

Nerocila munda Harger, 1873. Uncommon; an ectoparasite on various fish. 

Sphaeroma quadridentatum Say, 1818. Along shores, under stones, among algae, on 
peat banks. 

Suborder Valvifera 

Chiridotea almyra Bowman, 1955. In water of low salinity, as in the Pocasset Riv- 
er, along with Cyathura polita . 

Chiridotea coeca (Say, 1818) . Burrows like a little mole beneath the surface of 
intertidal sand flats. Common. 

Chiridotea nigrescens Wigley, 1961. Recently described from brackish waters of 

tidal marshes, North Falmouth to Chatham, on the southern side of the Cape. Re- 
sembles the more common C_. coeca , which is more marine in its habitat. 

Chiridotea tuf tsi (Stimpson, 1883) . On subtidal sand and mud bottoms in marine 
situations; sometimes concealed by mud adhering to rough back. 

Edotea triloba (Say, 1818) . On muddy shores, usually with adherent dirt. Edotea 
montosa (Stimpson, 1853) is probably a synonym (Wallace, N. A., 1919. Univ. 
Toronto Studies, Biol. Ser. no. 18: 42 pp.) 

Erichsonella attenuata (Harger, 1873). Apparently uncommon. 

Erichsonella f iliformis (Say, 1818) . Among eelgrass and algae. Common. 

Idotea balthica (Pallas, 1772). ifbiquitous. 

Idotea metallica Bosc, 1802. Common, swimming or clinging to vegetation. 

Idotea phosphorea Harger, 1873. Among weeds and on gravelly bottoms. 

Suborder Asellota 

Asellus sp. or spp.. Asellus is characteristic of fresh waters or of very low sal- 
inities. Although the name Asellus communis Say, 1818, has been the most widely 
used in the past (see, for example, Van Name, 1939) it now seems that Asellus 
militaris is the most common eastern American species (Van Name, 1942) . For 
specific determination, the assistance of a specialist is advised. 

J aera marina (Fabricius, 1780). Very common if looked for among weeds, mussels. 

Suborder Oniscoidea 

Ligia oceanica (Linnaeus, 1767) . Has been reported in numbers near high tide mark 
on rocky shores and under boards north of Boston. Its reported occurrence at 
Woods Hole needs confirmation. 

Phi l osc ia vittata Say, 1818. Found along the shore near high water mark, under 
boards and rubbish. 

S cyphacella ar enico la Smith, 1873. Found in the sand of beaches 

Note: See Van Name, 1936, for other terrestrial as well as fresh water isopods. 

Isopods 107 

Suborder Epicaridea (Bopyroidea) 

lone thomp_sqivi Richardson, 1904. On Calliana ssa; the type locality is North Fal- 
mouth . 

Probopyru s p andalicola (Packard, 1879) . On Palaemonetes . in the gill chamber. 

St egophryxus hypti us Thompson, 1902. Thompson reported 3-4% of hermit crabs Pag - 
uru s lonaicarpuB at Hadley Harbor infested, and 1.5% at Woods Hole. Parasite 
occurs on the abdomen of its host. 


Bayliff, W. H., 1938. A new isopod crustacean (Sphaeromidae) from Cold Spring Har- 
bor, Long Island. Trans. Amer. Micr. Soc, 5^.: 213-217. 
Bowman, T. E., 1955. The isopod genus Chiridotea Harger, with a description of a 

new species from brackish waters. J. Wash. Acad. Sci., 45_: 224-229. 
Menzies, R. J., 1957. The marine borer family Limnoriidae (Crustacea, Isopoda) . 

Part I. Northern and Central America: systematics, distribution, and ecology. 

Bull. Marine Sci. Gulf and Caribbean, 7: 101-200. 
Miller, M. A. amd W. D. Burbanck, 1961. Systematics and distribution of an es- 

tuarine isopod crustacean, Cyathura polita . Biol. Bull., 120 : 62-84. 
Richardson, H. , 1905. A monograph on the Isopods of North America. Bull. U. S. 

Nat. Mus., No. 54: 1-727. 
Richardson, H. , 1909. The isopod crustacean, Ancinus depressus (Say). Proc. U.S. 

Nat. Mus., 36. : 173-177. 
Swan, E. F., 1956. Isopods of the genus Ligia on the New England coast. Ecology, 

37: 204-206. 
Van Name, W. G. , 1939. The American land and fresh-water isopod Crustacea. Bull. 

Amer. Mus. Nat. Hist., 71: 1-535. 
Van Name, W. G., 1942. A second supplement to the American land and fresh-water 

isopod Crustacea. Ibid ., 80: 299-329. 
Wallace, N. A., 1919. The Isopoda of the Bay of Fundy. Univ. Toronto Studies, 

Biol. Ser. no. 18: 42 pp. 
Wigley, R. L. , 1961. A new isopod, Chiridotea nigrescens , from Cape Cod, Massa- 
chusetts. Crustaceana, 2_: 286-292. 


By Eric L. Mills 

Amphipods are a widespread and important group of crustaceans. That they are 
poorly known is mostly due to their wide limits of morphological variation, making 
taxonomy difficult, and partly due to lack of patience in investigators, since 
long and careful scrutiny is sometimes necessary. Since members of the group are 
found in a wide variety of ecological niches, they are of interest to students of 
taxonomy, ecology, physiology, genetics, and behavior. 


Amphipods have the basic malacostracan body plan of 6 head, 8 thoracic, and 
6 abdominal segments, but there appear to be only 7 free thoracic segments because 
of the fusion of the first thoracic segment with the head capsule. This first tho- 
racic segment bears the maxillipeds, and each of the 7 free thoracic segments bears 
a pair of limbs. The abdomen is divided into a pleon of 3 segments, each bearing 
a pair of pleopods and a urosome of three segments, each with a pair of uropods. 
A terminal telson projects from the rear of the urosome. The appendages are as 
follows : 


Plate 14 


Sources in Plates 14 and 15: Redrawn from Richard- 
son (R) , from Kunkel (K) , from specimens (S) , or 

as noted. 

Fig. 1. Tanais cavolini , with chela 
at greater scale (R) . 

3. L. savignyi , female (R) . 

5. Cirolana concharum (S) . 

7. Limnoria liqnorum , with details 
of pleotelson, after Menzies. 

9. Nerocila munda (R) . 

2. Leptochelia savignyi , male (R) . 

4. Cyathura polita (R) . 

5. Lironeca ovalis (K) • 

8. Limnoria tripunctata , details 
of pleotelson, after Menzies. 

10. Idotea baltica (S) . 

11. Same, ventral view of pleotelson 12. Idotea metallica (R) . 
with valve-like left uropod raised, 
exposing pleopods. 

13. Telson of Idotea phosphorea (R) 

15. Erichsonella f ilif ormis (R) . 

17. Asellus militaris (R) . 

19. Philoscia vittata (R) . 

14. Edotea triloba (R) . 

16. Telson of E. attenuata (R) . 

18. Jaera marina (R) . 

20. lone thompsoni , female in dor- 
sal view at left, male enlarged 
at right (S) . 

21. Probopyrus thompsoni , female at left with male beside it to same scale; 
male enlarged at right (R) . 

22. Steqophryxus hyptius , female in dorsal view at left, in ventral view 
at right with male indicated at same scale, male enlarged in center (R) 

Plate 14 


Plate 15 


Fig. 23, Chiridotea almyra ; A. head, B. first 
pereipod of male, C. pleotelson; redrawn 
after Bowman. 

24. Chiridotea tuftsi : A, B, C, as above. 

25. Chiridotea coeca : A, B, C, as above. 

26. Ancinus depressus (R) . 

27. Sphaeroma quadridentatum (R) . 

28. Exosphaeroma papillae , redrawn after Bayliff. 

29. Ligia oceanica (R) . 

30. Scyphacella arenicola (R) . 

31. Hyper i a ga lba , hyperiid amphipod (K) . 

32. Aeqinella lonqicornis , caprellid amphipod (K) 

33. Caprella geometrica , caprellid amphipod (K) . 

Plate 15 


23 b 


24 b 



23 c 


24 c 

25 c 


112 Amphipods 

1-2: Antennae - (two pairs) - both with peduncles and flagella, first antenna often 
with accessory flagellum. 

3; Mandibles - with molar and incisor surfaces and a palp. 

4: First maxillae - with inner and outer lobes and a 2-segmented palp. 

5: Second maxillae - simple, of inner and outer lobes. 

6: Maxillipeds - with inner and outer plates and a 4-segmented palp. 

7-13: Seven peraeopods, each 7-segmented, including flattened coxal plates which con- 
tribute to the sides of the body. The first 2 pairs of peraeopods are often called 
gnathopods and are subchelate, with the 6th segment expanded as a hand. When this 
"gnathopod 1 and 2" terminology is used, the remaining peraeopods are numbered from 
1-5. . 

14-16: Pleopods (3 pairs) - with a basal peduncle and 2 many jointed setose rami. 

17-19: Uropods (3 pairs) - with a basal peduncle and 2 more or less spinose single 
segmented rami. 

For illustrations, see Chevreux and Fage (1925, figs. 1 and 2). 

(After Kunkel, 1918) 

1. Eyes absent, but articulated eye lobes present INGOLFIELLIDEA 

1. Eyes present, sessile 2 

2. Eyes large, covering side of head; uropods with laminate 

rami, forming a tail fan with telson HYPERIIDEA 

2. Eyes not covering whole side of head 3 

3. Abdomen well developed GAMMARIDEA 

3. Abdomen rudimentary; body very slender (typical caprellids) 

or very much flattened ("whale-lice") CAPRELLIDEA 

The Hyperiidea are mostly pelagic; some are commensals with scyphozoan jelly- 
fish (Plate 15, fig. 31). Caprellidea (figs. 32, 33) are common inhabitants of 
weedy areas in littoral and sub-littoral zones. See Chevreux and Fage (1925) for 
common families and genera of these two suborders. Ingolf iellidea are rare, highly 
aberrant, mostly abyssal forms, unfamiliar to most taxonomists. Gammaridean amphi- 
pods are abundant and obvious members of all aquatic communities, but at present 
it seems advisable to present only a key to families. For figures, the references 
must be consulted. 

(Based on Stebbing, 1906) 

1. Antenna 1, first joint stout, with accessory flagellum; 
mandible cutting edge almost smooth; gnathopod 2 with 3rd 
joint elongated LYSIANASSIDAE 

1. These characters not combined 2 

2. Head tapering, truncate; eyes simple, usually 4; antennae 

1 without accessory flagellum; telson cleft AMPELISCIDAE 

2 . These characters not combined 3 

3. Peraeopods 3-5 with heavy armature of spines and setae, 

joints greatly expanded 4 

3. Peraeopods 3-5 less setose, joints not greatly expanded 5 

Amphipods 113 

4. Peraeopod 4 not greatly longer than peraeopod 5 HAUSTORIIDAE 

4. Peraeopod 4 greatly longer than peraeopod 5 PHOXOCEPHALIDAE 

5. Antenna 1 without accessory flagellum; maxillipeds abnormal; 

telson entire; fourth coxal plates greatly enlarged STENOTHOIDAE 

5. These characters not combined 6 

6. Mandible with molar weak or wanting; telson divided; maxilli- 

ped inner plate small LILJEBORGIIDAE 

6. These characters not combined 7 

7. Eyes dorsally continuous or confluent; pronounced rostrum; 

peraeopod 5 elongated OEDICEROTIDAE 

7. Eyes lateral, rostrum not well developed, peraeopod 5 not 

greatly elongated 8 

8. Side plate 4 excavate behind; peraeopods 1& 2 not glandu- 
lar; animal not tube dwelling 9 

8. Side plate 4 usually not excavate behind; peraeopods 1 & 2 

glandular; animal often tube dwelling 13 

9. Mandible with palp 10 

9. Mandible without palp 12 

10. Telson entire CALLIOPIIDAE 

10. Telson usually cleft, antenna 1 with accessory flagellum, 

urosome with patches of dorsal spines GAMMARIDAE 

10. Telson cleft; other features unlike previous combination 11 

11. Gnathopod 1 without hand BATEIDAE 

11. Gnathopods 1 & 2 without hands, telson cleft, antennae of 

male with calceoli PONTOGENEIIDAE 

12. Uropod 3, both rami well developed DEXAMINIDAE 

12. Uropod 3, one ramus small or wanting TALITRIDAE 

13. Pleon compressed 14 

13. Pleon usually depressed; tubicolous animals, often band- 
ed with color COROPHIIDAE 

14. Uropod 3 uncinate, i.e. with short hooks 15 

14. Uropod 3 not uncinate PHOTIDAE 

15. Inter-antennal lobes small; gnathopod 2 palm usually simple, 

hand not expanded AMPITHOIDAE 

15. Inter-antennal lobes prominent; gnathopod 2 palm often com- 
plex, hand expanded ISCHYROCERIDAE 

Animals may be most readily placed at least to genus by using the keys in Steb- 
bing (1906) and by reference to Sars (1895), Holmes (1905), Kunkel (1918) and Chev- 
reux and Fage (1925). Kunkel is particularly useful for local species. The follow- 
ing list of local genera and species is by no means complete but may aid in narrow- 
ing the search, and is presented merely as an aid to those starting a study. The 
development of keys to the species of amphipods in the Woods Hole region must await 
further studies. We are indebted to Dr. E. L. Bousfield for assistance in the com- 
pilation of the following list. 

114 Amphipods 



Lysianopsis alba Holmes, 1903. In sand, gravel or mud in shallow subtidal. 
Orchomenella pinguis (Boeck, 1861) . 
Tmetonyx nobilis Stimpson, 1853. 


Ampelisca macrocephala Liljeborg, 1852. 
Ampelisca vadorum Mills, 1963. 
Byblis serrata Smith, 1873. 


Found burrowing in sandy beaches. A complex family of several genera, current- 
ly being revised. 


Paraphoxus epistoma (Shoemaker, 1938) . 
Paraphoxus spinosus Holmes, 1903. 
P hoxocephalus holbolli (Kroyer, 1842). 


Stenothoe spp. 

Lil jeborgiidae 

Listriella clymenellae (Mills, 1961) . Commensal in tubes of Clymenella torquata . 


Monocu lodes edwardsi Holmes, 1903. 


Calliopius laeviusculus (Kroyer, 1838) . Commonly taken swimming at night lights. 


Carinogammarus mucronatus (Say, 1818). In both marine and brackish waters. 

Cr angonyx spp. In fresh water. 

Elasmopus laevis (Smith, 1873) . 

Gammarus annulatus Smith, 1873. Marine. 

Gammarus f asciatus Say, 1818. In fresh water. 

Gammarus oceanicus Segerstrale, 1947. Marine and brackish water. 

Gammarus tiqrinus Sexton, 1939. Brackish waters. 

Melita nitida Smith, 1873. 


Batea c atharinensis Fr. Muller, 1865. 


Pontogeneia inermis (Kroyer, 1838) . 



ripyami np the a Boeck, 1861. 



Allorchestes littoralis Stimpson, 1853. Intertidal. 
Hyale spp. Intertidal. 

Hvalella azteca (Sausseure, 1818). In fresh water. 
Orchestia spp. Rather small, dark, beach hoppers. 

Talorchestia longicornis (Say, 1818). The common, large whitish beach hopper. 
Talorchestia megalophthalma (Bate, 1862) . A beach hopper similar to the above but 
smaller and less common. 

Corophiidae (a family of tube dwellers) 

Cerapus tubular is Say, 1818. Among masses of Amaroucium , and among eelgrass. Said 

to carry tubes about. 
Corophium spp. In soft tubes. 
Eric thonius spp. 
Unciola irrorata Say, 1818. Often found in tubes of polychaetes or other amphipods. 


Leptocheirus pinguis (Stimpson, 1853) . 
Leptocheirus plumulosus Shoemaker, 1932. 
Photis spp. 


Ampithoe lonqimana Smith, 1873. 
AmpithQe rubricata (Montagu. 1808) 
Cymadusa compta (Smith, 1873) . 


Ischyrocerus spp. 

Jassa falcata (Montagu, 1808) . 

Chelura terebrans Philippi, 1839. A wood borer. 

Sympleustes qlaber (Boeck, 1861) . 


Lembos smithi (Holmes, 1905). 
Microdeutopus damnoniensis (Bate, 1856) . 
Microdeutopus gryllotalpa Costa, 1853. 


Barnard, J. L. , Index to the families, genera and species of Gammaridean Amphipoda 
(Crustacea). Allan Hancock Publications, Occas . Papers _19: 1-145. Very useful 
in literature searches. 

116 Amphipods 

Bousfield, E. L., 1963. Studies on amphipod crustaceans of the Cape Cod region. 

Preliminary report, Systematics-Ecology Program, MBL, Woods Hole (unpublished) . 
Chevreux, E., & L. Fage, 1925. Amphipodes, _in Faune de France, Paris, 9^: 1-488; 

438 text figures. Very useful for fauna of this region. Includes Hyperiidea 

and Caprellidea. 
Holmes, S. J., 1905. The Amphipoda of southern New England. Bull. Bur. Fisheries 

for 1904: 457-529, pi. 1-8. 
Kunkel, B. W. , 1918. The Arthostraca of Connecticut. Conn. Geol. and Nat. Hist. 

Survey 26^: 1-261. Standard work on Amphipoda of New England. 
Mills, E. L., 1962. A new species of Lil jeborgiid amphipod with notes on its bio- 
logy. Crustaceana, 4: 158-162. 
, 1963. A new species of Ampelisca (Crustacea: Amphipoda) from eastern 

North America, with notes on other species of the genus. Canad. J. Zool., 41 : 

Sars, G. 0., 1895. The Crustacea of Norway. Christiania & Copenhagen, _1: 1-711; 

240 pis. A standard reference, illustrations excellent. 
Stebbing, T. R. R., 1906. Amphipoda 1. Gammaridea. Das Tierreich (Berlin) 21 : 

1-806; 127 text figures. The basic reference work in amphipod taxonomy. 

Chapter XIII H 7 


Class Crustacea 

The Higher Malacostraca (Decapoda and Stomatopoda) 

The great majority of the well known crustaceans collected and studied at Woods Hole 
are in the order Decapoda, although the group also includes some little known forms and 
presents problems of identification in certain families, such as in the Xanthidae and 
Pinnotheridae . The "mantis shrimps" of the order Hoplocarida or Stomatopoda (see page 
127) are represented only by two species, not commonly collected. 

Figure references in this chapter are to Plates 16 and 17. 


I. Small animals of shrimp-like form; abdomen well developed, with tail fan; pleopods 
used for swimming; pleura (side plates) of 2nd abdominal segment overlapping those of 
first segment; the typical shrimps and prawns (fig. 41) Section CARIDEA 1 

II. Animals of heavier form, with well developed abdomen; pleopods not used for swim- 
ming; thoracic legs adapted for walking; chelipeds large and strong. 

A. Abdomen not much (if at all) wider or longer than cephalothorax; body firm, well 
armored and well pigmented; the familiar fresh water crayfishes and, as the only 
local marine representative, the lobster Homarus americanus 

B. Abdomen markedly wider and/or longer than cephalothorax; bodies soft, seemingly 
weak; burrowing forms with tendency to reduced eyes and pigmentation; sometimes 
placed with Anomura Subsection THALASSINIDEA 4 

III. Animals of typically crablike form, with 5th (last) pair of thoracic legs not 
markedly reduced; abdomen reduced, lacking uropods, and folded under body; antennae 
short and set medial to eyes; the "true crabs" Section BRACHYURA 5 

IV. Fifth pair of thoracic legs very reduced and folded up above bases of 4th pair; 
form varies from guite crablike to not very crablike; abdomen in some is asymmetrical 

or twisted, or reduced, but with uropods Section ANOMURA 27 

I. Caridea 

1. Carpus (the segment next to the claw or hand) of 2nd pair 

of legs not annulated (fig. 40) 2 

1. Carpus of 2nd pair of legs annulated into 3 parts (fig. 39); 

with humped abdomen (Family HIPPOLYTIDAE) Hippolyte zostericola 

2. Rostrum short, not compressed; first pair of legs subchelate 
(fig. 31); 2nd pair of legs chelate (as in fig. 41); eyes set 

close together (Family CRANGONIDAE) Crangon septemspinosus 

2. Rostrum long and laterally compressed; first and 2nd pair of 

legs chelate; eyes set widely apart (Family PALAEMONIDAE) .... Palaemonetes 

There are three local species, of which P. vulgaris is much the most common; the tabula- 
tion below should separate the three: 



Dorsal rostral 

Ventral rostral 

Dorsal rostral 
teeth lying post- 
erior to orbits 

Teeth of chelae 

of second legs : 

movable finger 

fixed finger 

P_. vulgaris 
(figs. 33. 34) 

to tip 


P_. intermedius 
(figs. 35. 36) 

to tip 


P. pugio 
(figs. 37. 38) 

tip naked 


II. Thalassinidea 

3. Chelipeds unequal in size; body very pale 

Callianassa atlantica 

3. Chelipeds of equal size 4 

4. Chelipeds with fingers not deflexed (fig. 30); fairly 

common and of good size (ca. 100 mm long) Upogebia af finis 

4. Chelipeds with fingers deflexed (subchelate, fig. 29) ; 

a rare animal of small size (to ca. 35 mm) Naushonia crangonoides 

Note: The Hoplocarida (Stomatopoda) or "mantis shrimps" have somewhat the same habitat 
and body build as the above 3 species, but are at once distinguished by their for- 
midable mantis-like claws, large eyes, and by the short carapace which does not cover 
the last 4 thoracic segments (see page 127 and fig. 32). 

III. Brachyura 

5. Carapace triangular or globose, with apex projecting for- 
ward to form a rostrum; "spider crabs" 6 

5. Carapace usually broader than long; rostrum small or want- 
ing 9 

6. Carapace triangular, resembling small chip of stone (fig. 3); 
walking legs very small but chelipeds relatively huge (Family 

PARTHENOPIDAE) Heterocrypta granulata 

6. Walking legs long and conspicuous 


7. Carapace with medial dorsal spines 8 

7. Carapace surface smooth, without median dorsal spines; splotched 

with red Pelia mutica 

8. With 6 spines in the median dorsal row; few dorsal tubercles 

(fig. 1) Libinia dubia 

8. With 9 spines in median dorsal row; many dorsal tubercles 

(fig. 2) Libinia emarginata 

Decapoda 119 

9. Free living crabs, well pigmented, with eyes not reduced 10 

9. Small crabs with reduced eyes; commonly but not always with 
reduced body pigmentation; usually commensal in worm tubes or 
bivalves, but some may wander free; carapaces either subcir- 
cular or markedly widened from side to side; small, rarely 

reaching 2.5 cm across carapace 19 

10. Carapace outline varies, but characteristically front mar- 
gin is curved, and bears a series of teeth between the eye 
and the (antero) lateral corner on each side; size varies, 
but several over 5 cm across carapace belong here; "can- 
croid" crabs 11 

10. Carapace outline squarish, with a more or less straight 
front margin; most are active, semi-terrestrial crabs, 
rarely over 4 cm across carapace; free living "grapsoid" 

crabs 23 

11. First antennae (antennules) folded longitudinally or nearly 

so (fig. 4) Family CANCRIDAE 12 

11. First antennae folded transversely or obliquely (fig. 5) 13 

12. Edges of antero-lateral teeth entire; chelipeds granulate, 

not denticulate (fig. 6) Cancer irroratus 

12. Edges of antero-lateral teeth denticulate; upper margin of 

palm denticulate (fig. 7) Cancer borealis 

13. Last pair of walking legs not markedly adapted for swimming; 

tips sharp, for walking (fig. 10) 14 

13. Last pair of legs flattened and paddle-like; tips rounded, 

for swimming (figs. 8, 9) 18 

14. Front (region between eyes) produced into 3 low teeth; 5 
very prominent sharp anterolateral teeth (fig. 10); hind 
pair of legs slightly flattened; an active long legged crab 
commonly 5-7.5 cm across carapace; the common "green crab" 

but color varies from greenish-black to orange Carcinus maenas 

14. Front not produced into teeth; generally small crabs, but 
more heavily built and less active than Carcinus , with shorter 
legs and heavier chelae; tips of chelae dark in some species; 

a group in which specific identification requires care 
Family XANTHIDAE 15 

15. Frontal margin (between eyes) with a single edge, not seem- 
ing double; fingers of chelae dark 16 

15. Frontal margin transversely grooved so as to appear double; 
fingers of chelae whitish; carapace with 4 and 3 distinct 
transverse lines of granules on anterior half 

Rhithropanopeus harrisi 

16. Movable finger of major cheliped with a heavy blunt tooth 
near base (fig. 26); carapace width often exceeds 2.5 cm 

(fig. 25) Panopeus herbsti 

16. Movable finger of major cheliped without such a tooth 17 

17. With an elongated dark red spot on the inner (concealed) 
surface of each maxilliped (note: Panopeus herbsti some- 
times but not always has a comparable spot, but it is 
small, circular, and located near base of maxilliped) ; fin- 
gers of minor chela "spooned" (fig. 28) ; carapace rather 
flattened and oval in outline (fig. 27) ; uncommon 

Eurypanopeus depressus 

17. Without such red spot on maxillipeds; fingers of minor chela 
not "spooned" (Fig. 24); carapace outline nore angular (fig. 
23) not oval; very common Neopanope texana 

120 Decapoda 

18. Antero-lateral teeth on carapace 3 to 5 in number (fig. 9); 

carapace not usually broad; color cream to tan with red mark- 
ings ("lady crab") Ovalipes ocellatus 

18. Antero-lateral teeth 9 in number (fig. 8); carapace extremely 
broad; outermost lateral tooth especially long and sharp; with 
some blue coloration, particularly on chelipeds ("blue crab") 

Callinectes sapidus 

19. Fifth (last) pair of thoracic legs very small and tucked up 
under carapace; you are in the wrong part of key; animal is 
a brachyurous anomuran ; go on to no. 30. 

19. Fifth (last) pair of thoracic legs is not especially reduced, 
useful in locomotion. This is a difficult family with marked 

sexual dimorphism Family PINNOTHERIDAE 20 

20. Carapace thin and shiny; body almost circular (fig. 12); com- 
mensal in mantle cavity of bivalves or in Chaetopterus tubes 

Pinnotheres maculatus 

20. Carapace much wider than long, generally commensal 

Pinnixa 21 

21. Propodus (2nd segment from tip) of 3rd walking leg slender, 
twice as long as wide (figs. 13, 14) 

Pinnixa sayana 

21. Propodus of 3rd walking leg not slender; less than twice as 

long as wide 22 

22. Body and legs smooth, not especially hairy; propodus of 3rd 
leg only slightly if at all longer than wide (figs. 15, 16) 

Pinnixa cylindrica 

22. Body and legs distinctly hairy; propodus of 3rd leg clearly 
longer than wide, and with edge granular or serrated (figs. 

17, 18) Pinnixa chaetopterana 

23. A gap is left between the 3rd maxillipeds when they are held 
at rest; front very or moderately broad; eyestalks short 

Family GRAPSIDAE 24 

23. The 3rd maxillipeds almost or quite close over the mouth region 
when held at rest; front moderately or very narrow; eyestalks 
long; one cheliped of male very large; local representatives 

are the typical "fiddler crabs" Family OCYPODIDAE 

24. Carapace very square (fig. 19); dark plum colored to bluish black 
(the 'square backed fiddler"); found burrowing in salt marshes 

Sesarma reticulatum 

24. Carapace with corners rounded off (fig. 20) ; color variably 
blotched olive-green; drifts in on Sargassum ('Columbus crab", 

"Gulf-weed crab") Planes minutus 

25. Inner surface of large claw of male with an oblique ridge 

(fig. 22) 26 

25. Inner surface of large claw of male without such oblique 
ridge (fig. 21) ; carapace mottled in grays and purple 
("calico back") Uca pugilator 

Decapoda 121 

26. Carapace uniformly dark, almost black; not exceeding as a 

rule 22 mm in width; very numerous (the "black fiddler crab') 
Uca pugnax 

26. Carapace grayish, lighter at front, with central brown 
mark; size very much larger than in the common fiddler crabs; 
carapace commonly over 23 mm, even 36 mm wide; reddish spots 
at joints of chelipeds, especially in males ("big fiddler 

crab" red- jointed fiddler crab") Uca minax 

IV. Anomura 

27. Animals inhabit gastropod shells; abdomen soft and twisted 

Chermit crabs") 28 

27. Abdomen symmetrical and tucked under thorax 30 

28. Chelipeds slender; hands subcylindrical 29 

28. Hands broad, flat, tuberculate Paqurus pollicaris 

29. Chelae with 4 or 5 distinct purplish bands on whitish back- 
ground Paqurus annulipes 

29. Chelae without such distinct banding Paqurus lonqicarpus 

30. Form essentially crablike, except for having 5th (last) 
thoracic legs reduced and hidden under carapace (fig. 11) ; 
small, rounded; chelipeds directed forward in resting posi- 
tion (not transversely as in Brachyura) ; antennae long and 
set lateral to eyes (Family PORCELLANIDAE) 

Polyonyx macrocheles 

30. Body egg shaped; carapace gray and shining; telson forms 
a long triangular white ventral shield; burrows in wave- 
swept sandy beaches (Family HIPPIDAE; "mole crabs", "sand 
bugs") Emerita talpoida 

I. Caridea (shrimps) 

Craqo , see Crangon . The re-establishment of the old (1798) generic name Crangon for the 
common shrimps is the result of the suppression in 1955 of the name Craqo by the In- 
ternational Commission on Zoological Nomenclature (Opinions and Declarations, 10( 1) : 
1-44) . This action terminated a most confusing situation, briefly: In 1904 Rathbun 
(Proc. Biol. Soc. Wash. 17 : 170) reported that the name Crangon had first been used 
in an obscure work by Weber (1795) for the "snapping shrimps" known since 1798 as 
Alpheus , and in consequence she redesignated Alpheus as Crangon . This left the com- 
mon shrimps without a name, so she adopted for them the "next available" name, Craqo . 
The remarkable 3 -way confusion included the use of the family name Crangonidae for 
the snapping shrimps and Cragonidae for the common shrimps. Under the 1955 ruling 
our common shrimp is secure under the generic name Crangon (Family Crangonidae) , and 
the snapping shrimps (which do not occur at Woods Hole) are Alpheus , and restored to 
the arms of their family Alpheidae. 

Crangon septemspinosus Say, 1818. Found in sand, in which it buries itself, below tide 
marks. Common, the name Crangon vulgaris Fabricius, 1798 has been used by those who 
believe the local form to be conspecific with the common European shrimp. 

Hippolyte zostericola (Smith, 1874) . Particularly common among eel grass ( Zostera ) , its 
green body color rendering the shrimp inconspicuous. 

Palaemonetes intermedius Holthius, 1949. Prior to 1949 this species and the' next were 
confused with and included in P. vulgaris . Found in brackish water, and mainly 









Plate 16 


Sources: Redrawn from Rathbun (R) , from 
Ryan (Ry) , from specimens (S), all but nos. 
13-14 by Bruce Shearer. 

. 1. Libinia dubia , young male, carapace (R) . 

Libinia emarginata , young male, carapace (R) . 

Heterocrypta granulata (R) . 

Lower side of front of Cancer irroratus to show first antennae folded 
longitudinally or obliquely (S) . 

Same region of Carcinus maenas to show first antennae folded transversely 

Carapace of Cancer irroratus , to show smooth edged anterolateral teeth (S) 

Anterolateral margin only of Cancer borealis , showing denticulate antero- 
lateral teeth (S) . 

Callinectes sapidus , carapace and fifth leg (S) . 

Ovalipes ocellatus , carapace and 10. Carcinus maenas , carapace and 
fifth leg (S) . fifth leg (S) . 

Polyonyx macrocheles (S) . 12. Pinnotheres maculatus , male (S) . 

Pinnixa sayana , male, ventral (S) . 14. Same, female (S) . 

Pinnixa cylindrica , male (S) . 

Pinnixa chaetopterana , male (S) . 

Sesarma reticulatum (S) . 

Inside of large claw of male Uca 
pugilator , lacking oblique ridge 

Carapace of Neopanope texana 

Carapace of Panopeus herbsti 
(points exaggerated) (S) . 

Carapace of Eurypanopeus depres - 
sus, to show oval outline (Ry) . 

16. Same, female (S) . 

18. Same, female (S) . 

20. Planes minutus (S) . 

22. Same, but of Uca pugnax , with 
oblique ridge (S) . 

24. Major chela of same, with area 

of black outlined extending onto 
palm (S) . 

26. Major chela of same, to show blunt 
tooth on movable finger (S) . 

28. Minor chela of same, to illustrate 
"spooned" condition (Ry) . 

Plate 16 


Plate 17 


Figs. 29-40: Redrawn from Kingsley (K) , Holthius 
(H) , specimens (S) , by Bruce Shearer. 

Fig. 29. Naushonia cranqonoides . carapace and left chela (K) . 

30. Upoqebia af finis {S\ left cheliped only. 

31. Crangon septemspinosus , "subchelate" first chela (S) . 

32. Squilla empusa (order Stomatopoda, not Decapoda) (S) . 

33. Second chela of Palaemonetes vulgaris (H) . 

34. Rostrum of same (S) . 

35. Second chela of Palaemonetes intermedius (H) . 

36. Rostrum of same (H) . 

37. Second chela of Palaemonetes pugio (H) . 

38. Rostrum of same (S) . 

39. Second leg of Hippo lyte to show annulated carpus (S) . 

40. Second leg of Palaemonetes ; carpus not annulated (S) . 

41. Lateral view of a generalized decapod, redrawn after Schmitt, 
showing the typical shrimp-like ("caridoid") form and append- 
ages . 

Plate 17 

126 Decapoda 

Palaemonetes pugio Holthius, 1949. In brackish to almost fresh water. 

Palaemonetes vulgaris (Say, 1818). This is the species usually collected and used at Woods 
Hole. Tends to occur in more saline water than the two less common species above. 

II A. Astacura or Nephropsidea 

(Crayfish and lobsters) 

Homarus americanus Milne-Edwards, 1837. The familiar lobster, generally taken commercial- 
ly in subtidal water; uncommon locally because of fishing. A point for comparative 
physiologists to bear in mind is that the lobster is more closely related to the fresh 
water crayfishes than to any other of the local marine crustaceans. 

II B. Thalassinidea 
(Mud shrimps) 

Callianassa atlantica (Smith, 1874) . A burrower in muddy sand, intertidal and in shallow 
water. The name C. atlantica replaces the name C. stimpsoni , which has prior appli- 
cation to a Pacific coast fossil species (see Schmitt, 1935) . 

Naushonia crangonoides Kingsley, 1895. Rare; has been found in intertidal or shallow 
subtidal, in muddy sand. 

Upogebia af finis (Say, 1817) . Habitat similar to that of Callianassa , but muddier. 

III. Brachyura 
(True crabs) 

Callinectes sapidus Rathbun, 1895. The blue crab. Common in grassy bays and salt water 
ponds. An important commercial and experimental animal, whose wide range on our east 
coast has been extended by accidental introduction into France, Holland, Denmark, and 
Israel. Penetrates into waters of very low salinity, especially in the warmer parts 
of its range. 

Cancer borealis Stimpson, 1859. Northern rock crab, Jonah crab. Despite name, is not 
confined to rocky areas but is often found in sand. 

Cancer irroratus Say, 1817. Rock crab. Similar to the above, but perhaps more charact- 
eristic of rocky areas . 

Carcinides , see Carcinus . " Carcinides " is now a rejected synonym of Carcinus . The facts 
are briefly: in 1897 Rathbun (Proc. Biol. Soc. Wash., 11 : 164) substituted Carcinides 
for the well known name, Carcinus , of the common green crab on the grounds that the 
name Carcinus had been given prior use by Latreille in 1796 for an essentially unrec- 
ognizable genus of amphipods . However, Latreille described no species under Carcinus , 
and the term in his sense was never subsequently used. In Opinion 330 (Opinions and 
Declarations, ... 9^(24): 321-338, 1955) the International Commission on Zoological 
Nomenclature suppressed Carcinus of Latreille 1796 and validated the Carcinus of 
Leach 1314 as applied to the green crab. 

Carcinus maenas (Linnaeus) . The green crab. The commonest crab of this area, in marine 
and brackish water. 

Eurypanopeus depressus (Smith , 1869) . Apparently uncommon or of doubtful occurrence in 

Woods Hole region; reported by Verrill and Smith but not taken by Sumner survey (1913) 
nor in the summer of 1963. It is extremely common further south (e.g. Delaware Bay), 
and is often associated with oysters or oyster shells in muddy areas where salinity 
may be lowered. 

Heterocrypta granulata (Gibbes, 1849) . Dredged on bottoms of shingles or pebbles, which 
the animal much resembles. 

Libinia dubia Milne Edwards, 1834. On muddy shores and flats. Stenohaline. 

Libinia emarginata Leach, 1815. Common on muddy shores and flats. This is the species 
more commonly used at Woods Hole. Stenohaline. 

Decapoda and Hoplocarida 127 

Neopanope texana sayi (Smith, 1869) . Extremely common. This subspecies ranges from 
Canada to Florida, where it merges and is confused with N. texana texana . 

Ovalipes ocellatus (Herbst, 1799) . Found in sand at low water mark and subtidally. 

Buries itself with only the eyes showing. Ranges from Cape Cod to South Carolina. 

Panopeus herbs ti Mi lne -Edwards , 1834. The species (as several "forms") ranges from 
Boston to Brazil. 

Pelia mutica (Gibbes, 1850) . Dredged, on gravelly and shelly bottom. 

Pinnixa chaetopterana Stimpson, 1859. Commensal in tubes of Chaetopterus , Amphi trite , 
and probably other worms. 

Pinnixa cylindrica (Say, 1818) . Said to occur in burrows of Arenicola crista ta . Young 
found free on low inter tidal muddy shores. 

Pinnixa sayana Stimpson, 1860. Reported to be dug out of mud, but probably actually 
in worm or other burrows. 

Pinnotheres maculatus Say, 1818. Commensal in Mytilu s , Modiolus , and several other bi- 
valves, as well as in Chaetopterus tubes. 

Planes minutus (Linnaeus) . A visitor that drifts into Woods Hole on Sarqassum ; may 
be found on shore occasionally. 

Rhi thr opanopeus harrissi (Gould, 1841) . This has never been reported from Woods Hole, 
but has probably been overlooked since its range is from Mexico to Maine and New 
Brunswick. It is found in waters of very low salinity, even fresh water, and has 
been introduced and become common in other regions, where it has been used in phy- 
siological research (e.g. San Francisco Bay, Holland, Germany) . 

Sesarma reticulatum (Say, 1817) . A southern form, of spotty ocurrence in certain fid- 
dler crab marshes, rather to landward. May be looked for in or near Uca minax hab- 

Uca minax (LeConte 1855) . A southern species, in this area near the extreme of its 
range and found only in a few localities. May occur in very low salinities. 

Uca pugilator (Bosc, 1801-02). In salt marshes; tends to make burrows in sand and in 
Salicornia areas. Less gregarious than U. puqnax . 

Uca puqnax (Smith, 1870). In salt marshes. Tends to make burrows in muddy areas. 

IV. Anomura 

Emerita talpoida (Say, 1817). Very common on exposed sandy beaches; intertidal. 

Pagurus annulipes Say, 1817. Dredged. 

Paqurus lonqicarpus Say, 1817. The commonest shallow water hermit crab; found everywhere. 

Pagurus pollicaris Say, 1817. The large hermit crab; common. 

Polyonvx macrocheles (Gibbes, 1850) . Lives in Chaetopterus tubes. 

Order Hoplocarida (Stomatopoda) 

(Mantis -shrimps) 

Members of this order differ greatly from decapods in structure, having only a 
short carapace that does not cover the last 3 or 4 segments of the thorax, and having 
the second maxillipeds modified to remarkable mantis-like, raptorial claws. They bur- 
row in subtidal or low intertidal mud, and are not commonly taken. There are two local 
species, distinguished as follows: 

1. Carapace and abdomen with several longitudinal carinae or keels; 
size large, up to 25 cm; 6 teeth on finger of raptorial claw; 
fairly common subtidally, but rare intertidally (fig. 32) 

Squilla empusa Say, 1818 

2. Carapace and abdomen smooth, without carinae; small, up to 
4 cm long; raptorial claw with 11-15 teeth; rare 

Nannosquilla grayi (Chace, 1958) 

(Described as Lysiosquilla grayi ; see Manning, 1963) 

128 Decapoda 


Chace, F. A., Jr., 1958. A new stomatopod crustacean of the genus Lysiosquilla from 
Cape Cod, Massachusetts. Biol. Bull., 114 : 141-145. 

Hay, W. P. and C. A. Shore, 1918. The decapod crustaceans of Beaufort, N. C, and the 
surrounding region. Bull. U. S. Bur. Fish. 35.(1915-1916): 371-475. 

Holthius, L. B., 1952. A general revision of the Palaemonidae (Crustacea, Decapoda, 

Natantia) of the Americas. II. The subfamily Palaemoninae. Allan Hancock Founda- 
tion Occ. Papers no. ^^2: 1-396. 

Kingsley, J. S., 1899. Synopsis of North American Invertebrates. III. The Caridea of 
North America. Amer. Nat. 3_3: 709-720. 

Manning, R. B. , 1963. Preliminary revision of the genera Pseudosquilla and Lysio - 
squilla . . . Bull. Mar. Sci. Gulf and Caribbean, J,3_: 308-328. 

Rathbun, M. J., 1917. The grapsoid crabs of America. Bull. U. S. Nat. Mus., 97 : 
i-xxii, 1-461. 

, 1925. The spider crabs of America. Ibid , 129 : i-xx, 1-613. 

, 1930. The cancroid crabs of America of the farailies Euryalidae, Port- 

unidae, Atelecyclidae, Cancridae, and Xanthidae. Ibid , 152 : i-xvi, 1-609. 

Schmitt, W. L., 1935. Mud shrimps of the Atlantic coast of North America. Smithson. 
Misc. Coll., 93: 1-21, 4 pi. 

Chapter XIV 129 

by W. Russell Hunter and Stephen C. Brown 

(Figure references in this chapter are to Plates 18 and 19 ) 

Among the relatively few patterns of animal construction (Phyla) which have 
been highly successful, in terms both of numbers and of diversity of habitats, is 
the molluscan plan. Along with the Chordata and the Nematoda, the Mollusca must 
rank just after the Arthropoda in terms of evolutionary "success". Most molluscs 
are readily recognizable — the soft body within a hard calcareous shell is usually 
diagnostic (as is the extensive use of ciliary and mucous mechanisms in feeding, 
locomotion, and reproduction) . The basic molluscan plan of structure and function 
is remarkably uniform throughout the group, but there is no single standard mollus- 
can shape. An extreme diversity in external body form has been based upon this 
plan — clam, chiton, snail, and squid are all molluscs. In an evolutionary sense, 
the molluscan plan has proved plastic, and the adaptive modifications of structure 
and function to fit particular environmental circumstances provide material for a 
wide variety of biological studies. 

There are probably about 90,000 living species of molluscs, mostly belonging 
to the three major classes, Gastropoda, Bivalvia, and Cephalopoda. In the mollus- 
can fauna of the Woods Hole area, the majority of orders and superfamilies are rep- 
resented, but students and visiting investigators should note that certain groups 
are absent and others represented by a single common species. For example, in the 
class Gastropoda, no two gilled aspidobranchs (Haliotidae, Fissurellidae, etc.) 
occur, and of the one gilled aspidobranchs only the limpet Acmaea testudinalis is 
available. There are no shallow water trochaceans (the group including Marqarites 
and Tegula ) such as are common on other coasts. Representatives of the class Sca- 
phopoda ("tusk shells", e.g. Dentalium ) can only be dredged from deeper water and 
are not available for experimental studies. Although most bivalve groups are well 
represented, including three typical genera of the subclass Protobranchia, the sub- 
class Septibranchia is not found locally. Among the Cephalopoda, neither octopods 
nor sepioids are available locally, though Octopus has been dredged from deeper 
water offshore. 

This chapter includes an outline classification of Mollusca, a glossary of 
terms relating particularly to shelled forms, some general references, a short sec- 
tion on local amphineurans and cephalopods, and detailed keys and annotated lists 
of species for the two large groups, the shelled gastropods and the bivalves. For 
nudibranchs and other shell-less ophistobranch gastropods, see Chapter 15. 

Much molluscan systematics in the past has been based on shell characteristics, 
and previously available keys have been almost entirely conchological - the vital 
soft parts and environmental circumstances being ignored. We have deliberately 
tried to design the present keys to be used on molluscs collected alive - and 
therefore they will not be so useful for the identification of stray empty shells 
found cast up on the beach. We feel, however, that by using such characters as the 
siphons of bivalves, we have created less artificial keys. We have tried to use 
criteria of more fundamental biological significance than the detailed dimensions 
of shells, and attempted to avoid the artificial separation of naturally related 
genera. In almost all cases, the specific names used follow those of Abbott (1954). 
Occasionally, where a well known generic name is no longer correctly applied to a 
particular species, we have given this after an "equals" sign in parentheses , e.g. 
( = Venus ) . These "former" genera should be clearly distinguished from the few sub- 
genera quoted, which are placed in parentheses following accepted usage. 

The outline classification given below is that which has been used in teaching 

130 Mollusca 

at Woods Hole for three years, and reflects modern reconsideration of phylogeny in 
the light of functional morphology (see, for example, the classification of the bi- 
valves) . Typical genera (local if possible) are given in parentheses, aid are marked 
"*" if species occur in the Cape Cod area, "n" if non-marine. 


A. Class MONOPLACOPHORA ( Neopilina) 


I. Subclass APLACOPHORA ( * ChaetodermaJ 

II. Subclass POLYPLACOPHORA (* Chaetopleura) 

C. Class SCAPHOPODA (* Dentalium) 



a. Order Archaeogastropoda 

(Diotocardia or Aspidobranchia) (* Acmaea ) 

b. Order Mesogastropoda 
(Monotocardia I or Taenioglossa) 

(* Littorina , * Lacuna , * Crepidula ) 

c. Order Neogastropoda 
(Monotocardia II or Stenoglossa) 

(* Busycon , * Nassarius , * Urosalpinx ) 

II. Subclass OPISTHOBRANCHIA (key to shell-less forms is in Chapter XV) 

a. Order Cephalaspidea (* Acteon , * Philine ) 

b. Order Anaspidea (* Haminoea , * Aplysia ) 

c. Order Thecosomata (Thecosomatous Pteropods) . . . . (* Cavolin a) 

d. Order Gymnosomata (Gymnosomatous Pteropods) (* Clione ) 

e. Order Sacoglossa (* Elysia ) 

f. Order Acochlidiacea ( Hedylopsis ) 

g. Order Notaspidea ( Pleurobranchus ) 

h. Order Acoela (Nudibranchia) . . . . ( * Acanthodoris , * Aeolidia ) 


a. Order Basommatophora (* Melampus , n* Lymnaea ) 

b. Order Stylommatophora (n* Cepaea ) 

E. Class BIVALVIA (Lamellibranchia or Pelecypoda) 

I. Subclass PROTOBRANCHIA (* Nucula , * Yoldia , * Solemya ) 

II. Subclass LAMELLIBRANCHIA (no longer used to designate whole class) 

a. Order Taxodonta (* Anadara ) 

b. Order Anisomyaria (* Mytilus , * Chlamys , * Crassostra ) 

c. Order Heterodonta (* Tellina , * Mercenaria ) 

d. Order Schizodonta (n* Anodonta , n* Unio ) 

e. Order Adapedonta ( * Mya , * Ensis ) 

f . Order Anomalodesmata (* Lyonsia , * Pandora) 

Mollusca 131 

III. Subclass SEPTIBRANCHIA ( Cuspidaris ) 



II. Subclass AMMONOIDEA (entirely extinct) 


a. Order Decapoda (* Loligo ) 

b. Order Octopoda (* Octopus ) 

c. Order Vampyromorpha ( Vampyroteuthis ) 


Adductor muscle ( s ) : The major muscles (usually two in number) of the bivalve body, 
inserting on each valve, the contraction of which closes the shell valves. 

Aspidobranch ctenidium: (Gastropods) A "shield" or "feather shaped" gill with 
leaflets or filaments alternating on either side of the axis (figs. 1, 2) 

Body whorl: The last and largest whorl which terminates at the aperture (Gastro- 
pods) (fig. 9) 

Byssus : (Bivalves) A permanent (or more usually temporary) attachment of tough or- 
ganic threads secreted from a gland in the foot. 

Callus: A local area of shell thickening ( a callus may occur near the umbilicus 
in certain snails) . 

Cardinal teeth: Hinge teeth immediately below the umbo. 

Chondrophore: (Bivalves) An internal shelf near the hinge line, often spoon shaped 
which bears the "resilium", or internal ligament in certain groups of clams. 

Columella: The thickened axis of the shell about which the whorls are developed 
(Gastropods) (fig. 9) . 

Concentric sculpture: Centering around the umbo and parallel to the shell margin 
(Bivalves) . 

Ctenidia (sing. - ctenidium): The characteristic gills, present and structurally 
homologous in all major groups of the Mollusca. 

Dimyarian: (Bivalves) With two approximately equal adductor muscles (figs. 5, 17). 

Exhalent siphon: (Bivalves) The more dorsal (nearer the hinge line) tubular ex- 
tension of the mantle edge through which water passes out from the mantle cav- 
ity, actually from the so-called "suprabranchial chambers" above and within the 
gill lamellae. 

Foot: The muscular locomotory organ in all molluscs. (In Cephalopoda it is rep- 
resented by the siphon and possibly the tentacles) . 

Growth lines or rings: (Bivalves) Lines concentric to the umbo on each valve which 
mark the successive positions of the shell margin during earlier growth. 

Heteromyarian : (Bivalves) With two markedly unequal adductor muscles (figs. 7, 16). 

Hinge teeth: See cardinal teeth, lateral teeth. 

Inhalent siphon: (Bivalves) The more ventral tubular extension of the mantle edge 
through which food bearing water is drawn into the mantle cavity. 

Inhalent siphon: (Gastropods) The gutter shaped extension of the thickened edge 

of the mantle through which water is drawn into the mantle cavity in some snails. 

Labial palps: Paired ciliated triangular flaps on either side of the mouth in bi- 
valves . 

Lamellar gills: Enlarged, flattened plate-like gills (ctenidia) which form the feed- 
ing organs of most bivalves (figs. 7, 8). 

Lateral teeth: Hinge teeth (anterior and posterior to the cardinals). 

Ligament: The elastic horny hinge of the bivalve shell. 

Longitudinal sculpture: Crossing the direction of growth of the whorls, essential- 
ly axial to the spiral (Coiled gastropods) . 

Lunules: (Bivalves) Heart shaped impressions (fig. 28 ) in the midline of the shell 
anterior to the umbones (Venerid clams) . 

132 Mollusca 

Monomyarian: (Bivalves) Apparently with only a single large adductor muscle 

uniting the valves (fig. 18). 
Operculum: A horny or calcareous plate attached dorsally to the posterior end 

of the foot in gastropods, which forms a trap door closing the aperture of 

the shell on withdrawal of the animal. 
Pallets: Paired flat shelly plates (figs . 25-2 7) secreted at the distal end of the 

elongate siphons in the worm-like wood borers (important in species syste- 

matics) . 
Palp proboscides: Long ciliated grooved tentacles extending from the labial 

palps (found only in protobranchiate bivalves) (fig. 5). 
Pectinibranch ctenidium: (Gastropods) A "comb shaped" gill with leaflets or 

filaments extending from one side of the gill axis only, the axis usually 

being fused to the "roof" of the mantle cavity (figs. 3, 4). 
Radula: A tough chitinous ribbon bearing teeth of various forms, part of the 

buccal apparatus in all molluscs except bivalves, used to obtain food by a 

rasping, "licking" action. 
Radial sculpture: Radiating from the umbo (Bivalves). 
Ribs: (Bivalves) Radial sculpture. 
Ribs: (Gastropods) Longitudinal sculpture. 
Spiral sculpture: Running from the shell apex to the aperture along the whorls 

(Coiled gastropods) . 
Siphonal canal, siphonal notch: A tube-like extension or notch-like infolding 

of the lip of the aperture in a gastropod shell through which the inhalent 

siphon is extended in life (figs. 9, 12). 
Sculpture: See concentric, longitudinal, radial, ribs, spiral. 

Taxodont dentition: A long row of many small uniform teeth in the hinge line (Bi- 
valves) (fig. 17) . 
Umbilicus: A pit or chink in the shell next to or within the base of the colum- 
ella, occurring in gastropods in which the largest whorls are not closely 

wound against each other axially(fig. 10). 
Umbo (pi. - umbones) : (Bivalves) The apparent "apex" or "beak" of each valve 

around which "radial" growth has proceeded. 


Abbott, R. Tucker, 1954. American Seashells 541 pp., Van Nostrand, Princeton. 

Abbott, R. Tucker, 1961. How to Know the American Marine Shells , 222 pp. Sig- 
net Key Books, New York. 

Fretter, V., and Graham, A., 1962. British Prosobranch Molluscs , their Function - 
al Anatomy and Ecology. 755 pp., Ray Society, London. 

Gould, A. A., and W. C. Binney, 1870. Report on the Invertebrata of Massachusetts, 
2nd Ed., vi + 524 pp., Boston. 

Heath, H. , 1918. Solenogastres from the eastern coast of North America. Mem. Mus. 
Comp. Zool. Harvard, 4_5: 183-261. 

Johnson, C. W. , 1934. List of marine Mollusca of the Atlantic Coast from Laborador 
to Texas. Proc. Bost. Soc. Nat. Hist., 40: 1-204. 

Miner, Roy Waldo, 1950. Field Book of Seashore Life , 888 pp., G. P. Putnam's 
Sons, New York. 

Morris, P. A. , 1947. A F ield Guide to the Shells of our Atlantic and Gulf Coasts 
236 pp., Houghton Mifflin, Boston. 

Morton, J. E., 1958. Molluscs , 232 pp., Hutchinson, London. 

Pelseneer, P., 1906. Mollusca , 355 pp. in "A Treatise on Zoology " (ed. E. R. 
Lankester) , A. and C. Black, London. 

Thiele, J., 1931-1935. Handbuch der systematischen Weichtierkunde 4 vols., Gus- 
tav Fischer, Jena. 

Verrill, A. E., 1882. Report on the cephalopods of the northeastern coast of Amer- 
ica. Rept. U. S. Fish. Comm, 1879. 

Verrill, A. E. and S. I. Smith, 1873. Report upon the Invertebrata of Vineyard 
Sound and Adjacent Waters, etc. Rept. U. S. Comm. Fish., 1871-72, pt. I: 
295-778, PI. 1-38. 

Mollusca 133 

Wilbur, K. M. and Yonge, C. M. , 1964. Physiology of Mollusca , 2 vols., Academic 

Press, New York. 
Yonge, C. M. , 1947. The pallial organs in the aspidobranch Gastropoda and their 

evolution throughout the Mollusca. Philos. Trans. Roy. Soc. London. B, 232 : 



Class Amphineura 
(Chitons and Solenogastres) 

The Polyplacophora (chitons) are more or less flattened animals bearing eight 
transverse calcareous plates dorsally. They possess a broad creeping foot, a well 
developed radula, and a variable number of paired ctenidia within the pallial grooves 
along each side of the foot. Chitons are typically found on hard substrates, such 
as the larger rocks of exposed shores or on shells in more protected waters. 

The Aplacophora (Solenogastres) are small shell-less forms that might not be 
recognized as molluscs by the general student. They have not been reported from the 
Woods Hole region proper, but a few species are known from deeper waters off the New 
England coast, the most common being Chaetoderma nitidulum Loven, 1845, from 50 me- 
ters or greater depths off Martha's Vineyard (Heath, 1918). 

In contrast to Solenogastres, chitons are well known to zoologists, and in some 
regions make up a conspicuous element of the intertidal fauna. The forty genera of 
Polyplacophora are, however, largely Indo-Pacif ic, and the only member of the group 
common at Woods Hole is: 

Chaetopleura apiculata (Say). "Common Eastern Chiton". Elongate-ovate; plates with 
15-20 rows of raised beads in the central area, with fine diagonal sculpture on 
the margins; tan or brown in color, girdle with reddish-brown to green mottling 
and covered with scattered short "hairs"; up to 3 cm in length; very common in 
shallow waters on stones and shells. 

A rarer species which has been reported taken from the gutters of Hadley Harbor is: 

Ischnochiton (= Trachydermon ) ruber (Linnaeus). "Northern Red Chiton". Elongate- 
ovate; surface of plates smooth except for growth lines, plates externally buff 
colored with reddish marblings, internally colored bright pink; girdle reddish- 
brown with minute elongate scales; up to 2.5 cm long; more common in deeper wa- 
ters north of the Cape. 

Class Cephalopoda 

The only local cephalopods are the squids. The common species, regularly avail- 
able for experimental work is: 

Loligo pealei Lesueur . "Common Squid" or "Long-finned Squid". Fins more than half 
the length of the trunk (fig. 30); without eyelids; color translucent gray with 
obvious chromatophores (mostly red and black) ; up to about 60 cm long; common, 
Cape Cod southward. 

Another squid occasionally but not regularly taken near Woods Hole is: 

Hex (= Ommastrephes ) illacebrosus (Lesueur) . "Northern", "Summer", or "Short-fin- 
ned Squid". Fins about V3 length of the trunk (fig. 31) ; with eyelids; color 
deep blue, ranging to red; up to about 45 cm long; common north of Cape Cod, 
but taken occasionally in Woods Hole waters in summer. 



1. Single aspidobranch ctenidium (figs. 1 and 2) and other arch- 
aeogastropod characters, simple conical shell without internal 
shelf or recurved apex, "tortoise-shell" coloring Acmaea testudinalis 

1. Single pectinibranch ctenidium (figs. 3 and 4), shells of var- 
ious forms -- coiled, conical, or reduced 3 

1. No ctenidium, mantle cavity as a lung, (and other Pulmonate 
characters), shiny ovoid shells with no operculum, small 

salt marsh or high littoral snails 2 

2. Squat, egg shaped, widest behind aperture, with very short 
conical spire, translucent brown, adult is about 15 mm long 

with 5-6 whorls, abundant in salt marshes Melampus bidentatus 

2. Top shaped, widest at aperture, with longer conical spire 
one-third length of shell, shiny brownish-yellow, adult is 
about 7.5 mm long with 7-8 whorls and incised suture, less 

common than Me lampus Ovatella .(^ Alexia ) myosotis 

3. Snails with an inhalent pallial siphon and an extensible pro- 
boscis bearing a narrow radula, mostly active predaceous car- 
nivores or carrion feeders, turbinate coiled shells with a 
canal or rudimentary notch next to the columella to accomodate 

the pallial siphon, (Neogastropoda - "Whelks" and "Drills") 31 

3. Snails other than above 4 

4. Shells with only apex as regular turbinate spiral shell, con- 
tinuing to grow as cylindrical tube which may be irregularly 

worm-like or minute cucumber shaped 7 

4. Cap shaped conical shells with internal shelf 5 

4. Snails other than above 9 

5. Almost circular "limpet" shell with central cup shaped shelf 

internally Crucibulum striatum 

5. Ovate "slipper limpet? shells with posterior shelf 6 

6. Robust "slipper limpet", shell opaque dirty white to tan, 
often with brownish blotches, shell height about one third 
of length, concave white shelf occupies about one half of 

aperture, often found in "stacks"; up to 5 cm long .... Crepidula fornicata 
6. Very flat "slipper limpet", always pearly white, variably 
flexed white shelf occludes less than one half of aperture, 
never forms "stacks", often inside shells or on Limulus ; 
up to 3.3 cm long Crepidula plana 

6. Small, relatively high "slipper limpet", apex obvious and 
often overhanging posterior margin of shell, oblique brown 
shelf occluding about one third of aperture; up to 13 mm long 

Crepidula convexa 

7. Attached tubular shell drawn out distally and irregularly 
twisted to resemble a large serpulid worm tube, relatively 
large, up to 7.6 cm long; only one species recorded near 

Woods Hole Vermicularia spirata 

7. Minute cucumber shaped molluscs with only a tiny spiral apex 

which may be eroded 8 

8. Glossy opaque white, with about 15 longitudinal ribs, about 

5 mm in length Caecum cooperi 

8. Translucent tan when alive, chalky white when dead, with about 

20-30 circular ribs; about 2 mm in length Caecum pulchellum 

Shelled Gastropoda 135 

9. Snails with no operculum, and reduced shell which is usually 

enclosed within expanded pallial or pedal tissues in life, 

animal usually incapable of withdrawing completely within 

shell 10 

9. Snails with normal operculum and shell, though expanded soft 

parts appear disproportionately large and may conceal shell 

almost completely; but animal is capable of total withdrawal 

into shell 15 

9. Snails with normal operculum and shell, soft parts are readily 

contained within shell 22 

10. Shell totally enclosed within animal 11 

10. Shell may be partially or largely exposed upon retraction of 

animal 12 

( Note : Pteropods and other pelagic gastropods with reduced 
shells are omitted from this key.) 

11. With internal saucer shaped horny shell, extensible mantle 
lobes used for swimming, the only species of "sea hare" 

recorded in this area; up to 20 cm long Aplysia willcoxi 

11. With fragile glassy bean shaped turbinate shell within man- 
tle, living animal characteristically exhibiting four fleshy 

lobes when viewed dor sally; up to 13 mm long Phi line lima 

12. Shell visible externally, with a prominent spire and ornamented 
with fine spiral rows of dots at level of aperture, shell up 

to 8 mm long Acteon punctostriatus 

12. Shells not usually visible externally when animals are active, 
spire low or absent, thin glassy shell lacking ornamentation, 

"Bubble-shell" Tectibranchs 13 

13. Shells with a small but elevated spire 14 

13. Shells with spire depressed into pit, that is, body whorl 

completely encloses the rest of the shell 15 

14. Stubby, fragile shell with very low spire, white with yel- 
lowish-brown staining, columella smooth; up to 3 mm shell 

length Retusa obtusa 

14. Moderately elongate, stronger shell with more obvious spire 
except when eroded; white with dark rust-brown staining; 
columella with strong spiral ridge; up to 6 mm shell length 

Retusa canaliculata 

15. Larger globose shell, fragile bluish-white; relatively large 

aperture; up to 13 mm shell length Haminoea solitaria 

15. Smaller elongate cylindrical shell; white with brown perio- 

stracum; relatively narrow aperture; up to 5 mm shell length . . Cylichna alba 
(Note: Other species of "Bubble-shell" Tectibranchs may occur 
in this area. ) 

16. Globose shells (shaped like Helicid land snails), lacking any 
siphonal canal on shell, with brown horny operculum character- 
istically D-shaped; in living animal enormous expanded foot 
partially encloses shell as animal plows along on sandy sub- 
strate; proboscis extrusible but not usually visible, short 
retractible fleshy inhalent pallial siphon; predaceous carni- 
vores (Naticid "Moon-shells") 17 

16. Other than above, recheck alternative characters at 3 and at 9. 

136 Shelled Gastropoda 

17. Shell umbilicus clearly open (fig. 10) 18 

17. Shell umbilicus totally or nearly occluded by callus 

(fig. 11) 20 

18. With no obvious callus 19 

18. With obvious ivory white thickened callus which does en- 
croach on the umbilicus; shell white with yellowish perio- 

stracum; up to 1 cm long Polinices immaculatus 

19. Coarse heavy shell, with no callus; brownish-gray; often 
with attached algal filaments; up to 12 cm shell diameter; 

common Lunatia heros 

19. Thin clean shell, with inconspicuous white callus on inner lip, 
light brown or white with th^ee characteristic rows of squar- 
ish dark brown spots on last whorl; up to 13 mm shell diame- 
ter; less common Lunatia triseriata 

20. Shell considerably wider than high with flat spire; clean 
bluish-gray, with obvious purple (or more rarely brown or 
pink) callus almost completely occluding umbilicus; up to 
7.5 cm diameter; often abundant inter tidally and in shallow 

W ater Polinices duplicatus 

20. Shells slightly higher than wide with obvious spire; white 
with pale brown periostracum; white callus closing or nearly 
closing umbilicus; opercula more calcified than other Nati- 

cids above; smaller species, less co..onon and usually subtidal 21 

21. Larger globose shell; polished white flat callus always com- 
pletely sealing over umbilicus; up to 3.8 cm shell height . . . Natica clausa 

21. Smaller ovate shell; white callus usually leaving open chink 
at umbilicus; shell usually with faint bands of light brown; 

up to 8 mm shell height Natica pusilla 

22. Elongate turret shells (fig. 15); usually more' than 5 obvious 

whorls; height more than 1.5 times diameter 23 

22. Globose shells (fig. 14); height less than 1.5 times diameter 29 

23. Shells coiled dextrally 24 

23. Shell coiled sinistrally; 10-12 whorls, dark brown with three 

spiral rows of prominent beads; up to 6 mm long Triphora niqrocincta 

24. About 11 markedly globose whorls, expanding rapidly to give a 
conical shell, each whorl bearing about 16 strong longitudinal 
ribs; circular aperture with thickened lip; up to 2.5 cm long; 

the only true "Wentletrap" recorded in this area Epitonium rupicola 

24. Whorls not markedly globose; whole shell awl or spindle shaped; 

mostly under 15 mm shell length 25 

25. Without obvious shell sculpture; about 5 somewhat globose 
whorls separated by a clear suture; aperture ovate-circular; 

up to 5 mm in length 26 

25. With obvious shell sculpture of various forms; more than 6 
somewhat flatter whorls; aperture flattened or rectangular; 

up to 18 mm long 27 

26. Minute; smooth yellow-brown shells with no markings apart 
from growth lines; up to 5 mm long (several species may occur 

in this area, including some in brackish waters) Hydrobia spp. 

26. Minute; light yellow to brown shell with microscopic spiral 
sculpture of incised lines; also tiny riblets near suture; 
up to 2.5 mm long Cingula aculeus 

Shelled Gastropoda 137 

27. Shell with rounded aperture with barely perceptible siphonal 
notch; 6-8 whorls when adult; about 5 nun long; common species 
which can be abundant intertidally Bittium alternatum 

27. Shells with obvious siphonal notch at anterior of aperture; 
10-15 whorls when adult; lengths 3 mm to 15 mm; rarer species, 

usually subtidal 28 

28. Minute species; glossy brown; later whorls bearing 2-3 rows 

of glassy beads; 10 whorls when adult; about 3 mm long. . Cerithiopsis greeni 
28. Larger species; more elongate; chocolate brown; flattish whorls 
each bearing 3 rows of distinct raised beads which are lighter 
in color; 14-15 whorls when adult; about 15 mm long . . Cerithiopsis subulata 

28. Larger species; more elongate; flattish whorls each bearing 

3 strong continuous squarish spiral cords; 10-12 whorls when 

adult; about 13 mm long (fig. 15) Seila adamsi 

29. Medium sized (up to 3 cm); coarser shells, lacking umbilicus 

or apertural groove; adults rarely translucent 30 

29. Very small, fragile, smooth shell with groove in inner lip 
extending into chink-like umbilicus; color variable from pink 

to brown with purple or dark brown markings ; up to 8 mm long . Lacuna vincta 
( Note : Minute specimens suspected of being spat (juveniles) 
of Littorina spp. or of Lacuna are best identified by comparing 
them closely with the apical parts of the shells in known spe- 
cimens of adults of all four species.) 

30. Spire flattened; shell very smooth and shiny; color variable 
but commonly uniform clear yellow-orange; often banded; up to 

1 cm across; in lower littoral associated with fucoid seaweeds 

Littorina obtusata 

30. Moderate spire, expanding more rapidly after first tiny whorls 

with raised spiral threads when young; usually black when young; 
variable but dull as adult; with planktonic larvae (thus apical 
shell is small and if not eroded in adult is very sharply point- 
ed) ; up to 3 cm across; thecommonest and most abundant "peri- 
winkle" at all tidal levels Littorina littorea 

30. Obvious but variable spire; shell rough with irregular raised 
lines; color variable but commonly greenish-yellow; viviparous 
(thus apical shell relatively large) ; up to 13 mm long; higher 

littoral (fig. 14) Littorina saxatilis 

31. Snails with protrusible proboscis and inhalent siphon, but with 
unsculptured globose shells lacking any siphonal canal or notch 

(almost certainly naticid "moon-shells") see 16 

31. Snails with protrusible proboscis and inhalent siphon, but with 
turreted shells usually with sculpture and with a siphonal canal 

or notch near the columella 32 

32. Minute turreted snails less than 5 mm shell length as adult 33 

32. Snails usually larger than 12 mm s>>ell length when adult (most 

species much larger) 34 

33. Smooth glossy shell; gray or yellow-brown with darkened marbl- 
ing; narrow oval aperture; with siphonal notch; up to 5 mm 
shell length, common free living species on eel grass and 

elsewhere in littoral Mitrella lunata 

33. White glossy shells with considerable ornamentation, usually 
no obvious siphonal notch; up to 5 mm shell length; usually 

ectoparasites (often on specific invertebrates) PYRAMIDELLIDACEA 

(See note overpage) 

138 Shelled Gastropoda 

Two species have been positively identified in this region: 
Odostomia ( Menestho ) bisuturalis - host unknown 
Odostomia ( Chrysallida ) seminuda - parasitic on Crepidula fornicata 

34. Siphonal canal short, forming conspicuous notch in apertural 

lip (fig. 13) 35 

34. Siphonal canal elongate, forming an obvious extension of the 

apertural lip (fig. 12) 39 

35. Larger "whelk"; solid chalky-gray shell, with rough yellowish 
periostracum; low but obvious sculpture consisting of about 5 
spiral cords crossed by about 12 longitudinal ribs in each 
whorl; flesh of living animal startling white with black blot- 
ches; up to 10 cm shell length; common offshore, rarely in 

littoral Buccinum undatum 

35. Smaller snails with conspicuous siphonal notch; less than 

2.5 cm long 36 

36. Stout "dirty" shell often eroded, dark brown or black with 
neither suture nor sculpture obvious; often with ad- 
herent debris or organisms; up to 2.5 cm long; often abundant 

on mud flats intertidally Nassarius obsoletus 

36. Clean shells with obvious sculpture 37 

37. Essentially conical shell with 8-9 whorls in adult, bearing 
about 5 spiral rows of ranked beads giving "waffle -like" pat- 
tern; off- white in color; up to 2 cm long(fig. 13); usually 

living on sand Nassarius trivittatus 

37. Sguat solid pear shaped shell with 4-5 whorls in adult, each 
bearing about 12 pronounced longitudinal folds; off-white in 
color; up to 13 mm long; less common in this area, living in 
muddy sand Nassarius vibex 

37. Slimmer spindle shaped shells, with 6-7 whorls in adult, with 

less complex sculpture; up to 18 mm long 38 

38. With about 12 low rounded longitudinal folds on each whorl; 

usually dark grayish-brown; up to 13 mm long; lower littoral . Anachis avara 

38. With about 24 narrow longitudinal folds on each whorl crossed 
by incised spiral lines; usually drab yellow in color; up to 

18 mm long; lower littoral and shallow water Anachis translirata 

( Note ; Specific identification of minute specimens suspected 
of being spat (juveniles) of Nassarius spp. or Anachis spp. 
is aided by comparing them with the apical parts of the shell 
in known - and not eroded - specimens of all five species) . 

39. Smaller "drills" etc. under 4 cm shell length as adults (most- 
ly about 2.5 cm long) ; common in littoral 40 

39. Larger "whelks" all over 6 cm length as adults (may be up to 

23 cm long) ; mostly sublittoral and offshore 43 

40. Relatively thin shelled with 6-8 globose whorls in adult; gray 
with greenish periostracum; no obvious shell sculpture; siphon- 
al canal of moderate length bent back from aperture; up to 2.5 

cm long Colus pygmaea 

40. Stout shelled "drills" with obvious shell sculpture 41 

41. With 5 angular whorls and deep suture, giving sharp "shoulders" 
to spire, and relatively long siphonal canal which is almost 

closed over; up to 2.5 cm long (fig. 12) Eupleura caudata 

41. With rounded but ornamented whorls, siphonal canal relatively 

short and open 42 

Shelled Gastropoda 139 

42. Very thick polished white shell with thickened lip (rarer color 
variants - clear yellow, orange -brownish or striped) ; about 5 
whorls in adult with rounded spiral ridges giving corrugated ap- 
pearance; up to 3.5 cm long; less common, on exposed rocky shores 
Thais ( = Nucella) lapillus 

42. Thick duller grayish, yellowish, or brown shell; about 6 whorls 
in adult, each with 9 to 12 strong longitudinal ribs crossed 
by spiral grooves giving knobby appearance; up to 2.5 cm long; 

the commonest local "drill," on all hard substrates .... Urosalpinx cinerea 
( Note : Minute specimens suspected of being spat (juveniles) of 
Eupleura , Thais , or Urosalpinx can best be separated by the rela- 
tive lengths of their siphonal canals. Comparison with apical 
parts of the shells of known adults is less useful here) . 

43. Combined length of aperture and siphonal canal about one half 

shell length 44 

43. Combined length of aperture and siphonal canal markedly greater 

than one half shell length 46 

44. Heavy grayish shell with 6-7 whorls as adult, bearing up to 
10 conspicuous reddish-brown spiral ridges; up to 10 cm long 
Neptunea decemcostata 

44. Lighter shells with simple globose whorls 45 

45. Shells more elongate, sharper spired; 6-7 whorls in adult; 
siphonal canal straight; thin semi -glossy periostracum; up to 

13 cm long Colus stimpsoni 

45. Shells less elongate, blunter spire; 5-6 whorls in adult; siphon- 
al canal usually twisted away from aperture; hairy periostracum; 

up to 6.5 cm shell length Colus pubescens 

46. Suture channeled giving broad flat "shoulders" to whorls; up 

to 20 cm long Busycon canaliculatum 

46. Suture not deeply incised; single row of knobby tubercles on 

inclined "shoulders" of whorls; up to 23 cm long Busycon carica 


Acmaea testudinalis (Muller) . "Tortoise-shell limpet". On hard substrata only, in 
the lower littoral and immediate sublittoral of exposed shores, the only true 
limpet in this area, and the single local gastropod species which retains the 
primitive feather shaped ctenidium of the rchaeogastropods . 

Acteon punctostriatus (C. B. Adams) . Near low water mark and offshore, preferred 
substrata not known. 

Anachis avara (Say). Lower littoral, common in some places. Both species of Ana - 
chis are predaceous carnivores and are more abundant in the south. 

Anachis translirata (Ravenel) . Lower littoral, but more commonly dredged in shal- 
low water . 

Aplysia willcoxi Heilprin. "Sea Hare". Probably rare, this is the only large 

tectibranch whih has been recorded in this area. The group is better repre- 
sented on more southern Atlantic coasts. 

Bittium alternatum (Say) . Common and often extremely abundant on tidal flats and 
in shallow water offshore, on a variety of substrata but especially organic 
muds arid eelgrass. 

Buccinum undatum L. "White helk" or "Buckie". Not uncommon offshore in colder 
waters, rarely in littoral, on a variety of substrata. 

(Note: *- Busycotypus canaliculars ; **- Busycon aruanum ; Hollister, S. C, 1958. 
Paleont. Amer. 4: 59-125) 

140 Shelled Gastropoda 

Busycon canaliculatum (L.) . "Channeled Whelk". Dredged in shallow water on sandy 
bottoms, more common than B. carica in this area. 

Busycon carica (Gmelin) . "Knobbed Whelk". Dredged in shallow water on sandy bot- 
toms , not uncommon . 

Caecum cooperi S. Smith, and 

Caecum pulchellum Stimpson. Minute cucumber shaped gastropods found interstitial- 
ly in sand or in dead sponges, detailed ecology unknown, inshore in warmer 

Cerithiopsis qreeni (C. B. Adams), and 

Cerithiopsis subulata (Montagu) . Less common in Cape area than in warmer waters 
to the south, in shallow water, detailed ecology unknown. 

Cingula aculeus Gould. In shallow water; other species of the Rissoacea may be 
discovered in this area. 

Colus pubescens (Verrill) and 

Colus pvgmaea (Gould) and 

Colus stimpsoni (Morch) . Dredged subtidally and offshore. 

Crepidula convexa Say. Littoral and offshore, the least common of the three local 
"slipper-limpets", occasionally on eelgrass and more commonly on a variety 
of shells. 

Crepidula fornicata (L.). "Common Slipper-limpet". Littoral and offshore, the com- 
monest and largest local "slipper-limpet", often found in stacks, showing char- 
acteristic sex change with size and position. 

Crepidula plana Say. Littoral and offshore, never forming stacks, variably flexed 
to fit their substratum which may be the inside surface of a larger molluscan 
shell, or the exoskeleton of Limulus . 

Crucibulum striatum (Say) . Dredged in shallow water, with an enlarged ctenidium 
like the closely related Crepidula spp. , probably also a filter feeder. 

Cylichna alba (Brown) . Low water mark and shallow water, the detailed ecology of 
this and other species of "Bubble-shell tectibranchs" is unknown. 

Epitonium rupicola Kurtz. "Brown-banded Wentletrap" . Near low water mark and fur- 
ther offshore, not uncommon, this is the only local species of a genus more 
common to the south. 

Eupleura caudata (Say). "Thick-lipped Drill". Hard substrata, lower littoral and 
subtidal, probably more common in warmer waters. This is the least common of 
the three local species of "drills" which are predaceous carnivores with radu- 
la and proboscis modified for boring and an accessory boring organ in the foot. 

Haminoea solitaria (Say). Low water mark and shallow water, probably the commonest 
"Bubble-shell tectibranch" in this area, may occur mainly in muddier inlets 
with eelgrass, detailed ecology and reproductive behavior unknown. 

Hydrobia spp. Several species of this difficult but widespread genus may occur in 
this area, some in brackish waters. 

Lacuna vincta (Montagu). "Chink-shell". Lower littoral and shallow water, often 
abundant, a cold water species. 

Littorina littorea (L. ) . "Common Periwinkle". Typically at midlevel littoral, in . 
this area the largest, commonest, and most abundant "periwinkle" at all tide 
levels, with planktonic larvae and less capacity to resist desiccation and to 
respire in air than L. saxatilis . 

Littorina obtusata (L.). "Smooth Periwinkle". At lower levels at the littoral, us- 
ually associated with fucoid seaweeds including Ascophyllum . This periwinkle 
is referred to in modern European physiological literature as L. littoralis 
(L.) (not to be confused with the distinct species L. littorea which also occurs 
in Europe), and in early U. S. conchological literature as L. palliata (Say). 

Littorina saxatilis (Olivi) . "Rough Periwinkle". Typically high level littoral, 
of variable shell form and color, in this periwinkle females have a brood 
pouch and give birth to shelled young resembling miniature adults, the gill 
is reduced and the mantle cavity more lung-like, and all stages have a great- 
er capacity to resist dessication than the other littorinids. L. rudis (Maton) 
is simply a synonym of L. saxatilis . 

Lunatia heros (Say). "Moon-shell". Not uncommon on sand intertidally but more abun- 
dant subtidally, this is the largest of the six local species of predaceous 

Shelled Gastropoda 141 

carnivores belonging to the Naticidae. It is less resistant than Polinices du - 
plicata to higher temperatures and lower salinities. 

Lunatia triseriata (Say). Less common and usually subtidal, this small naticid 
occurs, however, intertidally in Lagoon Pond, Martha's Vineyard, and elsewhere 
in this area. 

Melampus bidentatus (Say) (- M. lineatus (Say)). Usually living around the high wa- 
ter mark of spring tides, and abundant in the high levels of Cape Cod salt mar- 
shes, Melampus is a pulmonate belonging to the family Ellobiidae and thus prob- 
ably related to the gastropod stock which gave rise to most land snails and to 
the freshwater pulmonates like Lymnaea . 

Mitrella lunata (Say) . Near low water mark and in shallow water on a variety of 
substrata, often extremely abundant, this minute snail is closely related to 
Anachis spp. and is a predaceous carnivore using its extensible rasping and 
sucking proboscis to prey on various small sessile invertebrates such as Bot - 
ryllus . 

Nassarius obsoletus (Say). "Common Mud Snail". In shallow water subtidally, in- 
tertidally on mud flats, and even in estuarine conditions, this is often an ex- 
tremely abundant species congregating in packed masses which can temporarily 
cover acres. Embryologists have tended to refer to this snail as Ilyanassa 
obsoleta (using its subgeneric name) , and certain older books place it in the 
genus Nassa . 

Nassarius trivittatus (Say). Intertidally and in shallow water, usually on clean 
sand in sheltered localities, this species is common but less abundant than N. 
obsoletus . 

Nassarius vibex (Say) . Occurs on sand and mud flats intertidally, less common in 
this area than the other two Nassarius spp. 

Natica clausa Broderip and Sowerby. Dredged subtidally and offshore in the Cape 
Cod area, interti^al on some more northern shores. 

Natica pusilla Say. Dredged subtidally and offshore. 

Neptunea decemcostata (Say) . Dredged offshore, shells occasionally washed up. 

Odostomia (Menestho ) bisuturalis (Say) . Identified here, see below. 

Odostomia ( Chrysallida ) seminuda (Say) . Identified here, and found by the writers 
to live as an ectoparasite on Crepidula fornicata . 

( Note : Other species of the PYRAMIDELLIDACEA will probably be found in this 
area: they are usually ectoparasites and often highly specific as regards their 
host invertebrates.) 

Ovate 11a (= Alexia ) myosotis (Draparnaud) . Living around the high water mark of 

spring tides (for example, in crevices of wharfs and docks, and under the "trash 
line" of high salt marshes) , this minute pulmonate snail is related to Melam - 
pus . 

Phi line lima Brown. Not common in this area, dredged in colder waters. 

Polinices duplicatus (Say). "Shark Eye" or 'Moon-shell" . Most abundant intertidal- 
ly on sand, this is the commonest of the local species of the Naticidae. Unlike 
Lunatia heros , it survives lowered salinity and high temperatures, and is thus 
able to colonize estuarine and high littoral habitats. All naticid snails are 
predaceous carnivores using the radula in the extensible proboscis to drill 
through the shells of other molluscs. 

Polinices immaculatus (Totten) . Less common naticid, subtidal in this area, occur- 
ing more frequently to the south. 

Pteropods (and other pelagic gastropods with reduced shells) are omitted from this 

Retusa canaliculata (Say) . Sand and mud flats intertidally and in shallow water, 
this minute species plows along the surface like a tiny naticid and is prob- 
ably the commonest shelled tectibranch in this area. 

Retusa obtusa (Montagu) . Not uncommon in habitats similar to those of R. canali - 
culata . 

Seila adamsi (H. C. Lea). Not uncommon around low water mark and subtidally; like 
the related Cerithiopsis spp., this small "turret-shell" is less common in the 
Cape area than in warmer water to the south. 


Plate 18 


Figures 1-8: Mantle cavity and ctenidium in represent- 
ative molluscs. Figures 1-9 by Stephen C. Brown; 10-15 
by Ruth L. von Arx. 

Fig. 1. The aspidobrancb ctenidium as in the limpet, Acmaea testud - 
inalis . 

2. Face view of ctenidial leaflets in Acmaea . 

3. The pectinibranch ctenidium as in Littorina littorea . 

4. Face view of ctenidial leaflet in Littorina . 

5. The protobranch ctenidium, associated with large labial palps 
and palp proboscides, as in Nucula . 

6. Face view of ctenidial leaflets in Nucula . 

7. The true lamellibranch ctenidium, as in Mytilus . 

8. Face view of ctenidial filaments in Mytilus . 

9. Sectional view of the shell of BUCCJ nl1TTI undafcmi , showing whorls 
surrounding central columella from apical whorl at top to largest 
"body" whorl with prominent siphonal notch. 

10. The naticid "Moon-shell", Lunatia heros , showing the shell umbi- 
licus clearly open. 

11. The naticid "Moon-shell", Polinices duplicatus , showing the obvious 
pigmented callus nearly occluding the umbilicus. 

12. Eupleura caudata, showing the elongate siphonal canal forming an 
obvious extension of the apertural lip. 

13. Nassarius trivittatus , showing the short siphonal canal forming a 
prominent notch in the apertural lip. 

14. Littorina saxatilis , showing a typical "globose" shell with height 
less than 1.5 times diameter. 

15. Seila adamsi . showing a typical elongate "turret-shell" with height 
more than 1.5 times diameter. 

• Plate 18 



Plate 19 


Figures 16-20 and 25-29 by Mrs. R. von Arx; 
21-24 by W. Russell Hunter; 30-31 by Bruce 
Shearer . 

Fig. 16. Modiolus demissus , shell valve showing the heteromyarian 

condition of the adductor muscles, and a long external ligament. 

17. Anadara transversa , shell valve showing dimyarian condition of 
the adductor muscles, and taxodont dentition in the hinge line. 

18. Aequipecten irradians , shell valve showing monomyarian condition, 
of adductor muscle, and internal triangular ligament or "resiliuin" 

19. Thyasira gouldi , dorsal view of shell valves with characteristic 
prominent radial folds running posteriorly from anterior umbones. 

20. Macoma tenta . dorsal view of shell valves with characteristic 
posterior twist. 

21-24 (following). Siphonal types, and natural posture in substrate, 
of various bivalves. 

21. My a arenaria , a sedentary deep burrowing bivalve with massive fused 
siphons . 

22. Tellina aqilis , a deposit feeding bivalve with separate extensible 
siphons . 

23. Mercenaria mercenaria , an active shallow burrowing bivalve with 
short fringed siphons and a relatively massive foot. 

24. Ensis directus , a typical razor clam with short fringed siphons 
and massive powerful foot emerging terminally like a "mushroom" 
anchor . 

25. Bankia gouldi , a typical "Shipworm" bivalve; the elongate body 
bears reduced valves anteriorly and calcareous pallets at the si- 
phonal openings. 

26. Bankia gouldi , plume like compound pallet. 

27. Teredo navalis . simple paddle shaped pallet. 

28. Mercenaria mercenaria . anterior view of shell showing obvious 
deeply incised lunules below umbones. 

29. Pitar morrhuana . anterior view of shell showing large but indis- 
tinct lunules below umbones. 

30. Loligo pealei, showing characteristic lateral fins more than half 
the length of the trunk and large eyes without eyelids. 

31. I lex illa c ebrosus , showing characteristic short lateral fins and 
eyes with eyelids. 

Plate 19 



Thais (= Nucella ) lapillu s (L.). "Dogwinkle". Hard substrata, lower littoral and sub- 
tidal, common only in the Cape area on certain beaches exposed to wave action. 
This is another carnivorous "drill", feeding on barnacles as well as molluscs. 

Triphora nigrocincta (C. B. Adams). Around low water mark on seaweeds, not uncom- 
mon, this is the only local gastropod species with a sinistrally coiled shell. 

Urosalpinx cinere a (Say). "Oyster-drill". Hard substrata, lower littoral and sub- 
tidal, often abundant, this is the commonest of the three local species of drills 
but, besides using radula, proboscis, and pedal accessory boring organ to bore 
through the shells of other molluscs, this predaceous carnivore also feeds ex- 
tensively on barnacles, using muscular action by the proboscis to force apart 
the plates of the barnacle test without actual boring. 

Vermicularia spirata (Philippi) . Probably living in shallow water, detailed ecolo- 
gy unknown. "Worm-shells" of this type are rare in this area. Tucker Abbott 
(1954) records this species only for "Southeast Florida and the West Indies", 
but shells have been found in the Cape Cod area on several occasions and liv- 
ing specimens found once in 1961 and twice in 1963. 



1. Without lamellar gills, ctenidia resembling those of aspido- 

branch gastropods (figs. 5 and 6) Order Protobranchia 2 

1. With lamellar gills clearly modified for filter feeding (figs. 

7 and 8) 4 

2. Protobranchiates feeding by palp proboscides, without lamellar 

gills, with taxodont dentition (fig. 5) 3 

2. Modified protobranchiate with reduced palps, hinge of elongate 
shell without dentition, with lustrous yellow-brown periostra- 

cum, radially marked and extending beyond shell margins Solemya velum 

3. Obliquely ovate globose shell, usually greenish-gray Nucula proxima 

3. Asymmetric elongate shell, narrowing posteriorly, usually green- 
ish-brown Yoldia lima tula 

4. With lamellar gills, but retaining taxodont dentition (fig. 17) 5 

4. With lamellar gills, with other than taxodont dentition 6 

5. Rhomboidal shell (up to 3.8 cm); 30-50 ribs; gray-brown perio- 
stracum usually worn, longer stouter ligament clearly distinct, 

left valve overlapping right valve Anadara transversa 

5. Ovate shell (up to 5.9 cm); 26-35 ribs; hairy black-brown perio- 

stracum persistant; narrow ligament less distinct Anadara ovalis 

6. Monomyarian condition of adductor muscle (fig. 18) 7 

6. Markedly heteromyarian condition of adductor muscles (fig. 16) 12 

6. Dimyarian condition (or nearly so) of adductor muscles (fig. 

17) 19 

7. Swimming monomyarian bivalves (Scallops) 10 

7. Attached monomyarian bivalves (Oysters and Jingle shells) 8 

8. Lower bivalve with hole through which passes calcified byssus 9 

8. No such hole, lower valve directly cemented to hard substrata 

(Eastern oyster"/ Crassostrea virginica 

9. Upper valve translucent and smooth, up to 5.1 cm Anomia simplex 

9. Upper valve drably opaque and rough with small prickles . . . Anomia acu l eata 

Bivalvia 147 

10. Both valves with strong radial sculpture 11 

10. Rough, dirty-white, pink or yellowish valves without strong 

radial sculpture; up to 20 cm diameter Placopecten magellanicus 

11. Approximately equal 'Wings" (or 'ears") at hinge; 17-20 regu- 
lar ribs forming strong radial corrugations; up to 7.6 cm 

diameter (fig. 18) Aeguipecten irradians 

11. Markedly unequal "wings", about 50 irregular cord-like ribs; 

up to 10.2 cm diameter Chi amy s islandica 

12. "Shipworms" boring in timber, worm-like bodies (fig. 25) 

bearing reduced valves anteriorly and pallets posteriorly 18 

12. Wedge shaped shells, markedly narrow anteriorly with well 
developed, usually dark periostracum, byssal attachment, 
lacking siphons (Mussels) 13 

12. Markedly elongate shells, active burrowers in sand, with well 

developed foot and short or medium length separate siphons 16 

13. Umbo near but not at anterior tip of each valve (fig. 16) 14 

13. Umbo at anterior tip of each valve, no shell sculpture, blue- 
black with shiny periostracum; common in littoral, often abun- 
dant Mytilus edulis 

14. With radial shell sculpture and thin periostracum 15 

14. Without radial shell sculpture, with thick hairy periostra- 
cum; usually dark brown; up to 15 cm, less common species; 

lower littoral and subtidally Modiolus modiolus 

15. Hinge without teeth; strong radial ribs which bifurcate usu- 
ally covering entire shell; up to 10 cm long; salt marshes 

and upper littoral, often abundant (fig. 16) Modiolus demissus 

15. Hinge finely dentate, ribs on anterior and posterior thirds 
of shell only; up to 6.4 cm long; rarer species; subtidal in 

this area Musculus (- Modiolaria ) niger 

16. Umbo nearly central, siphons separate and moderately long; 

powerful foot emerging ventrally 17 

16. Umbo at anterior end of dorsal margin; very elongate shell 
curving dorsally; short siphons; powerful foot emerging 
terminally; "razor" shape; up to 25 cm long (fig. 24) Ensis directus 

16. Umbo towards anterior end; elongate ovate shell; short siphons; 
with strong internal rib running ventrally from umbo; up to 

6.4 cm long Siliqua costata 

17. Dull greenish-yellow periostracum often eroded around umbones ; 

thick opaque shell gaping at ends; up to 10 cm long Tagelus plebeius 

17. Shiny thin periostracum with radial markings; often pale pur- 
ple, fragile shell; up to 3.8 cm long Tagelus divisus 

18. Pallets as simple paddle shaped blades (fig. 27) Teredo navalis 

18. Pallets each plume-like of many cones (fig. 26) 
probably Bank i a gouldi (but check other Bankia spp.) 

19. Deep burrowing or boring bivalves with massive fused siphons 

(fig. 21) 20 

19. Deposit feeding bivalves with separate extensible siphons 

(fig. 22) 23 

19. Otherwise, usually with short fringed siphons (fig. 23) 29 

148 Bivalvia 

20. Stout rasping spines externally on anterior of valves, re- 
duced periostracum, borers in peat, clay, or stone 21 

20. Thin valves with irregular growth rings but no sculpture; 
thick periostracum on ventral margin and siphons; up to 15 

cm shell length; burrowing in muddy sand Mya arenaria 

21. Elongate shell, umbones nearer anterior, entire shell sculp- 
tured with spines; large siphons 22 

21. Squat shell; up to 5 cm length; umbones nearer midline; 
sculpture limited to anterior; shell gaping widely at both 

ends; disproportionately large siphons Zirfaea crispata 

22. Shell tapering posteriorly, with only slight gape; up to 

21 cm long Cyrtopleura ( =Barnea) costata 

22. Shell truncate posteriorly, gaping widely at both ends; up 

to 5 cm in length Cyrtopleura ( =Barnea) truncata 

23. Ligament internal, triangular 24 

23. Ligament external 25 

24. Symmetrical ovate shell; pearly white; up to 3.8 cm length 
Periploma ( z Cochlodesma ) leanum 

24. Asymmetrical, shell tapering posteriorly, chalky white; up 

to 2 cm in length Cumingia tellinoide s 

25. Shells markedly elongate 28 

25. Shells sub-circular or ovate 26 

26. Duller white shells, often with slight posterior twist (fig. 

20) , and pallial sinus more extensive in right valve 27 

26. Glossy white opalescent shell with pallial sinuses similar 

in both valves; up to 13 mm in length Tellina agilis 

27. Variable dull white shell, never elongate; up to 3.8 cm 

length Macoma balthica 

27. White, fragile shell moderately elongate-ovate; with slight 

but obvious posterior twist; up to 2 cm length Macoma tenta 

28. Shell sculptured with rasping spines; reduced periostracum; 
dull white borer in peat or clay; shell proportions somewhat 

variable Petricola pholadif ormis 

28. Shells not sculptured; well -developed periostracum 17 

29. Subg lobular shells 30 

29. Elongate ovate shell, tapering and becoming compressed post- 
eriorly; periostracum with fine radial lines, often with ad- 
herent sand grains; up to 2 cm shell length Lyonsia hyalina 

29. Entire shell compressed; with umbones near anterior corner of 
nearly rectangular valves; somewhat saddle shaped; up to 3.4 

cm across Pandora gouldiana 

30. Shells with concentric sculpture 31 

30. Shell with strong radial sculpture Cardita borealis 

30. Shells with no sculpture apart from irregular growth rings 34 

31. Ligament external 32 

31. Ligament internal, triangular; up to 10 mm across 

Cras s inella (=Gouldia) mactracea 

Bivalvia 149 

32. With lunules anterior to umbones (fig. 28) 

young Mercenaria (' Venus ) mercenaria 

32. Without such lunules 33 

33. Ovate with strong concentric sculpture; up to 5 cm across 

Astarte borealis 

33. Trigonal with low concentric sculpture; up to 2.5 cm across 

Astarte castanea 

34. Ligament external 39 

34. Ligament internal, triangular 36 

34. Ligament in groove but partly external; characteristic small 
shells with 2 prominent radial folds (fig. 19) running post- 
eriorly from umbones which are anterior 35 

35. Oblong valves, translucent-white; up to 13 mm long . . . Thyasira trisinuata 

35. Rounded valves, yellowish periostracum; up to 6.4 mm long . . Thyasira qouldi 

36. Mantle almost completely fused; short siphons surrounded by 
common circlet of tentacles; posterior of right valve slight- 
ly overlapping left; up to 6.4 mm long Corbula contracta 

36. Ventral margins of mantle apparently fused, but without tis- 
sue union; fourth pallial opening ventral to inhalent siphon 

with associated groove for waste disposal (mactrid clams) 37 

37. Mactrid clams less than 18 mm shell length 38 

37. Mactrid clam more than 18 mm shell length; strong, ovate; 
smooth shell covered by shiny thin yellowish-brown perio- 
stracum; spoon shaped chondrophore in left valve; up to 

18 cm shell length Spisula solidissima 

38. Rather globose shell with proportionately small chondrophore; 
no denticles on hinge teeth; each valve with a single low 

radial rib near posterior Mulinia lateralis 

38. Slimmer shells, with disproportionately large chondrophore; 
tiny saw-tooth denticles on anterior and lateral hinge teeth 
young of Spisula solidissima 

39. Shells under 5 mm shell length 40 

39. Shells over 5 mm shell length 42 

40. Look out for spat of larger bivalves ' 

a) If heteromyaria n with wedge shaped shell, specimen is likely to 

be a juvenile mussel. Go back to 13 

b) If moderately elongate with fused siphons, specimen is like- 
ly to be a juvenile of My a arenaria ; see 20 

c) If round or ovate with separate extensible siphons, specimen 

is likely to be a juvenile deposit feeder like a tellinid clam. 

Go back to 2 3 

d) If bearing prominent concentric sculpture and lunules anterior 
to umbones, specimen is likely to be juvenile of Mercenaria ; 

see 32 and 43 

40. Shells under 5 mm shell length, seemingly adult and not in above 

categories 41 

41. Minute globose shiny shell, with umbones near mid-line and 

fine concentric lines; white, often with blue or purple patches 

(very common and often abundant) Gemma gemma 

41. Minute ovate shell with posterior umbones and shiny nut-brown 
periostracum (rare and probably commensal) 

Mysella (= Rochef ortia ) planulata 

(See note overpage) 

2^50 Bivalvia 

(But other minute leptonacean clams may also belong here, check 
Aligena elevata and Montacuta = pp. ) 

42. Globular smooth "cockle" with variable brown patterning on 

white exterior of shell; interior always translucent yellow; 
actively moving through sandy substrates by large very exten- 
sible foot; up to 2.5 mm diameter Laevicardium mortoni 

42. Slimmer but still globose clams without above shell coloring; 

may be up to 15 cm in diameter 43 

43. Shell valves with marked, deeply incised lunules anterior to 
umbones (fig. 28) ; shell may retain traces of juvenile concen- 
tric sculpture, internal free margin of shells crenulated and 
usually purple; up to 15 cm diameter, "Quahog" 
Mercenaria (= Venus ) mercenaria 

43. Shell valves without such lunules and with shiny dark brown or 
black periostracum showing many fine incised growth lines; no 
crenulation on valve margin; up to 13 cm diameter 
Arctica ( - Cyprina ) islandica 

43. Shell valves with large but shallow lunules; dull chalky white; 
without crenulation on ventral margin of valves; up to 5 cm dia- 
meter (fig. 29) Pitar morrhuana 


Aequipecten irradians (Lamarck). "Bay Scallop". The commonest scallop of the area, 
in most larger harbors and lagoons on flats exposed at lowest tides, also sub- 
tidal in shallow water. 

Anadara ovalis (Bruguiere) . "Blood Ark" . Formerly known as Area campechiensis 
Gmelin, fairly common low water mark to subtidal. 

Anadara transversa (Say). "Transverse Ark". Smallest of our "Ark-shells" , fairly 
common in mud, low water mark to subtidal. 

Anomia acule^ta Gmelin. "Prickly Jingle". Less common, attached to rocks or emp- 
ty shells, near low water mark. 

Anomia simplex O rbiqnv. "Common Jingle" or "Mermaid's Toenail". More common, at- 
tached to logs, docks and boats as well as to rocks; near low water mark. 

Arctica (=Cyprina) islandica (L. ) ."Black Quahog". Dredged in muddy sand, common in 
certain areas. 

Astarte borealis Schumacher and 

Astarte castanea (Say) . Dredged in shallow water locally, may occur intertidally 
on exposed sand beaches north of the Cape. 

Bankia qouldi Bartsch. A common and very destructive "Shipworm" on the southern 
Atlantic coast, transported to this area in drifting wood. 

Barnea . see Cyrtopleura . 

Cardita borealis (Conrad). Dredged in shallow water and on the continental shelf, 
probably common in some places. This species is listed in most of the text- 
books as Venericardia borealis , but Venericardia is apparently a Tertiary 
genus not surviving today. 

Cerastoderma pinnulatum (Conrad) . Not in key. Dredged subtidally. 

C hlamys islandica Muller. "Iceland Scallop". Common in shallow water and on the 
continental shelf. 

Corbula contracta Say. Dredged in shallo'-' water. 

Crassinella (- Gouldia ) mactracea Lindsley. Dredged from current swept sand and 
shell bottom in shallow water. 

Crassostrea virginica (Gmelin). "Eastern Oyster". This is the commercially im- 
portant oyster of the Atlantic seaboard; greatly variable in size and shape; 
around and below low water mark in estuarine as well as marine conditions, 
attached to any hard substrata including empty shells, with free swimming 
larvae, unlike Ostrea spp. 

Bivalvia 151 

Cumingia tellinoides Conrad. In muds, usually associated with eelgrass, a deposit- 
feeding bivalve, which, however, does not belong to the family Tellinidae but 
to the Semelidae, illustrating convergence in functional morpholouy. 

Cyrtodaria siliqua (Spengler). Dredged subtidally. Not in key. 

Cvrtopleura (= Barnea ) costata (L.) . "Angel Wing". Around low water mark in sandy 
mud, may also be found boring in peat or clay. 

Cvrtopleura (- Barnea ) truncata (Say). "Truncated Borer". Borers in peat or clay 
on the Cape, and in softer rocks (e.g. shales) outside of this area. 

Ensis directu s (Conrad). "Common Razor Clam". Rapid vertical burrower in sand, 
very common in some areas (including Barnstable) . 

Gemma gemma (Totten) . Very common and particularly abundant on tidal flats. 

Hiatella gallicana (Lamarck) and possibly also 

Hiatella arctica (L.). Non-boring specimens belonging to this genus have been dred- 
ged subtidally and a living specimen of H. gallicana was found in 1963 in shal- 
low water at the Sandwich end of the Cape Cod Canal. Not in key. 

Laevicardium mortoni (Conrad). In muddy sand, occasionally with eelgrass, in more 
sheltered shallow waters. 

Lyons i a hyalina (Conrad) . Rare at low water mark, more commonly dredged. 

Macoma balthica (L.). Common in organic muds, a deposit feeder. 

Macoma tenta (Say). In muddy sand, a deposit feeder. 

Mercenaria (=Venus) mercenaria (L.). "Quahog" or "Little-neck Clam" or "Hard-shelled 
Clam". The most important commercial clam, very common, loyally abundant. 

Modiolus demissus (Dillwyn) . "Ribbed Mussel". Salt marshes and upper littoral, 
often abundant. The nomenclature, and even the systematic position, of this 
mussel have been obscure. Its shell lacks teeth next to the ligament, and it 
is certainly erroneous to place it in the genus Brachidontes , the shells of 
which always have tooth-like crenulations just behind the ligament. Care must 
be taken to distinguish M. demissus from Brachidontes recurvus Rafinesque 
("Hooked Mussel"), common on more southern Atlantic shores, but which may ex- 
tend into this area. Note that Mytilus (or Modiolus ) plicatulus is simply a 
synonym for M. demissus and that both Volsella and Arcuatula have been sugges- 
ted as generic names for this species. Volsella has been rejected, but recent- 
ly Soot-Ryen (1955) has suggested Arcuatula demissa as the best name after 
setting up several diagnostic characters (including the radial sculpture) to 
separate Arcuatula from the more characteristic species of the genus Modiolus . 

Modiolus modiolus (L.). "Horse Mussel". Cooler waters, usually below low water 
mark, reaching greatest size subtidally. 

Mulinia lateralis (Say) . A dwarf mactrid clam, related to Spisula , usually found 
in shallow water in mud or clay, but occasionally in the same habitat as Spi - 
sula in surf stirred sand. 

Musculus (= Modiolaria) niqer (Gray) . A rarer mussel, near low water mark and sub- 
tidally, more actively moving than most mussels. 

My a arenaria L. "Soft-shelled Clam" or "Long-neck Clam". The important commercial 
clam of mud flats, but occurring commonly in a wide variety of substrata (gra- 
vel, sand, mud or peat) at all tidal levels and in shallow water. Burrows are 
probably permanent and adult My a completely sedentary organisms. 

Mvsella (= Rochefortia ) planulata (Stimpson) . A minute leptonacean clam which is 
probably commensal. This is the only species of this group positively iden- 
tified recently, but Aligena elevata and Montacuta spp. may occur. Mysella 
planulata has previously been placed in the genera Kellia and Rochefortia . 

Mytilus edulis L. "Common Mussel". Common, often abundant in extensive, packed 

colonies forming 'blankets" , between tidemarks, and on the masonry and pilings 
of docks and other structures. 

Nucula proxima (Say) . In mud subtidally (several other Nucula spp. may occur in 
this area). Deposit feeding using palp proboscides, this is probably the 
most primitive genus of living bivalves. 

Pandor a qou l diana (Dall) . Rare, lower intertidal and shallow water. 

Periploma (=C ochlodesma ) leanum (Conrad) . Rare, dredged. 

Petricola phola diformis Lamarck. "False Angel Wing". The commonest norer in peat 
and clay in this area. This species, as is clearly revealed by the separate 
extensible siphons, is a modified deposit feeder derived from a stock like 

152 Bivalvia 

the Tellinidae, which shows secondary convergence in structure and habit with 
the Pholadidae, e.g. Cyrtopleura spp. and Zirfaea spp. 

Phacoides ( Lucinoma ) f ilosus Stimpson. Subtidally, dredged in Vineyard Sound. Not 
in key. 

Pi tar morrhuana (Linsley) . Dredged. 

Placopecten maqellanicus (Gmelin) . "Deep-sea Scallop". Common subtidally and on 
the Continental Shelf. 

Siligua costata (Say). On shallow water and sand flats , another vertically bur- 
rowing razor clam, belonging to the Solenidae. 

Solemya velum Say. Not uncommon on intertidal mud flats and mud subtidally, a spe- 
cialized representative of the Protobranchia. 

Spisula solidissima (Dillwyn) . "Surf Clam" or "Hen Clam". In surf stirred clean 
sand, common below low water mark on exposed ocean beaches, young specimens 
occur in the littoral. Called Mactra solidissima in much experimental literature. 

Tagelus divisus (Spengler) and 

Tagelus plebeius (Solander) (?T. gibbus (Spengler)). In muddy sand, on intertidal 
flats and in shallow water, these are deposit feeding bivalves related to the 
Tellinidae (note the separate extensible siphons) . Tagelus spp. have second- 
arily assumed structure and habits like those of the true razor clams. 

Tellina agilis Stimpson (=T. tenera Say) . In fine or muddy sand, on intertidal 
flats and in shallow water, the only local representative of this important 
world wide genus of deposit feeding bivalves with separate, very extensible 
siphons . 

Teredo navalis L. "Shipworm" . A common and destructive wood borer of world wide 
distribution, transported to this area in drifting wood from more southern 
parts of the Atlantic coast. 

Thyasira gouldi (Philippi) and 

Thyasira trisinuata (Orbigny) . Dredged. 

Yoldia l imatula (Say) . Dredged in shallow water just below low water mark, occa- 
sionally found intertidally (two other species, Yoldia sapotilla (Gould) and 
Yoldia thraciaeformis Storer, may occur in this area. This is one of the 
three local genera of Protobranchia, deposit feeding by means of palp probos- 
cides . 

Zirfaea crispata (L.). "Piddock". Borer in peat or clay on the Cape, and in soft- 
er rocks (e_.g_. shales) outside this area; more common in colder waters. 

Venericardia, see Cardita. 

Chapter XV 153 


Shell-less Opisthobranchia 

by George M. Moore 

Nudibranchs are the best known of the shell-less opisthobranchiate snails. Thiele 
(1931) recognized four orders of the subclass Opisthobranchia, placing some of the 
shell-less forms, along with a few shelled forms, in the order Sacoglossa. Most of 
the shell-less forms, along with a few shelled forms, were placed in the order Acoela, 
in two suborders, Notaspidea and Nudibranchia. Odhner (1934, 1936, 1939) recognized 
seven orders of Opisthobranchia including Notaspidea and Nudibranchia, and the class- 
ification adopted in Chapter XXV of this manual gives eight orders. The shell-less 
opisthobranchs of the Woods Hole fauna belong to three of these orders, with the ma- 
jority of them in the order Nudibranchia. 

Since the time of Alder and Hancock (1845-55) two general types of Nudibranchia 
have been recognized. One, the Holohepatica (dorids) , have the digestive gland (the 
so-called liver) compact and undivided. The other, the Cladohepatica (eolids) , have 
the digestive gland branched, with the branches extending into special dorsal out- 
growths of the mantle (the cerata) . Odhner proposed the division of the order Nudi- 
branchia into four suborders, since it appeared to him that the evolution of the 
branched digestive gland had occurred more than once. Three of the suborders are 
represented in this key and list. 

The suborder Doridacea is characterized principally by a circle of branchial 
plumes (adaptive or secondary gills) about the anus, which is mid-dorsal on the post- 
erior half of the animal. The digestive gland is compact and does not extend into 
dorsal extensions of the mantle. In some dorids the mantle is broad and distinct 
from the foot while in others it is reduced to a ridge or narrow fold. A single 
pair of tentacles is present, on the dorsal surface a short distance back from the 
anterior end. These are termed dorsal tentacles or rhinophores, in contrast to the 
oral tentacles found in certain other nudibranchs. In dorids these dorsal tentacles 
are delicately sculptured with several diagonal laminae. Typical dorids are repre- 
sented in the Woods Hole fauna by Palio , Onchidoris , and Acanthodoris . The family 
Corambidae, although included with the Doridacea, is not typical since its members 
lack the circlet of branchial plumes. They have a pair of small ctenidia-like gills 
located one on each side of the anus at the posterior end between mantle and foot. 

In the suborder Eolidacea branchial plumes are lacking and the anus is lateral- 
ly placed on the right side of the body. Typically, in all except the genus Emble- 
tonia , there are two pair of tentacles. The branched digestive gland extends into 
special dorsal processes (cerata; sing, ceras). Many eolids feed on hydroids and 
sea anemones, and undischarged nematocysts are moved by way of the "liver" canals 
to the cerata and concentrated in cnidosacs at their tips. A pore connects each 
cnidosac to the exterior. Eolids are represented in the Woods Hole fauna by nine 

The suborder Dendronotacea is represented in the Woods Hole fauna by Dendrono - 
tus, Scyllaea , and Idulia. In this suborder the dorsal tentacles are eguipped with 
a basal sheath into which the terminal club can be retracted. Both types of diges- 
tive gland organization are found among the genera of this suborder; the holohepa- 
tic condition is considered the more primitive. 

Nudibranchs, unless carefully anesthetized, contract and distort badly on pre- 
servation. Such specimens can be used for radula studies and for certain details of 
internal anatomy. This key will be of little use when working with such specimens, 
since it is intended for tentative identification of living specimens and is based 
entirely on external characteristics. Figure references in this chapter are to 
Plates 20 to 22. 


1. With branchial plumes arranged in circle about the anus; 
anus located medially on the posterior dorsal half of the 
body; with single pair of tentacles „ 2 

1. Without circlet of branchial plumes as above; other dorsal 
processes (pallial outgrowths) present or absent; with one 

or two pairs of tentacles 5 

2. Branchial plumes singly pinnate; mantle distinct from foot 3 

2. Branchial plumes doubly pinnate (or imperfectly tripinnate) ; 

mantle distinct or reduced to a ridge 4 

3. Body white or occasionally slightly yellowish; branchial 
plumes about 11 and set in circle close to anus; dorsal sur- 
face covered with numerous knobbed tubercles of variable 

size Onchidoris aspera 

3. Body dull yellowish -white (sometimes gray) with numerous 
brown markings; brownish markings form three indistinct inter- 
rupted longitudinal bands; branchial plumes up to 20-30 in 
large specimens, set in elliptical groove around anus, and 
with an included space covered by several bluntly knobbed 
tubercles; dorsal surface covered with numerous knobbed tu- 
bercles of variable size Onchidoris fusca 

4. Mantle distinct from foot; dorsal surface thickly covered 
with numerous soft, slender, conical papillae of almost uni- 
form size (figs. 3-5) Acanthodoris pilosa 

4. Mantle reduced to knobbed ridges extending from sides of head 
to alongside the circlet of branchiae; body sparsely covered 

with short, blunt tubercles of variable size Palio lessonii 

5. Cerata (dorsal processes which contain as a central core a 

branch of the digestive gland) absent 6 

5. Cerata present 10 

6. With distinct mantle; 1 or 2 ctenidia between mantle and foot 7 

6. Without distinct mantle; ctenidia absent 8 

7. With a single typical gastropod ctenidium on right side between 
mantle and foot; mantle covers anterior 2/3 of body but not the 

head (figs. 1, 2) Pleurobranchaea tarda 

7. With pair of small ctenidia between mantle and foot at posterior 
end of body in the midline; mantle covers entire body, including 

head (figs. 6, 7) Corambella (?) sp. 

8. Body compressed; with 2- pairs of strap-like membranous extensions 
arising from dorso-lateral edges of body; inner surface of dorso- 
lateral folds and surface between them thickly covered with deli- 
cately branched filaments (these filaments may be termed cerata 

by some authors) (fig. 8) Scyllaea pelaqica 

8. Body depressed; body with lateral extensions which can be folded 
towards midline over dorsal surface of body to enclose a dorsal 

canal 9 

9. Lateral folds about 2/3 of length of body, not extending to post- 
erior tip of foot, and not quite meeting when folded over back; 

foot square in front, with rounded corners; length to 1 cm . . . Elysia catula 
9. Lateral folds extending to posterior tip of foot, and overlapping 
each other when folded over the back; anterior angles of foot 
strongly extended with acutely angled tips; length to 3 cm 
Elysia chlorotica 

Shell-less Opisthobranchs 155 

10. Cerata branched and tree-like Dendronotus frondosus 

10. Cerata unbranched 11 

11. Single pair of tentacles (oral tentacles lacking) 12 

11. Two pair of tentacles (both oral and dorsal tentacles present) 13 

12. Tentacles arising from trumpet-like sheaths; cerata club 

shaped and bearing scattered dark colored tubercles .... Idulia corona ta 

12. Tentacles arising directly from head (without basal sheaths); 

cerata simple and without tubercles Embletonia fuscata 

13. Anterior lateral corners of foot extended and sharply acute 

angled 14 

13. Anterior lateral corners of foot not much extended and 

either rounded or bluntly angled 18 

14. Dorsal tentacles ringed with alternately larger and small- 
er rings, producing serrate margins (figs. 24, 25) . . Facelina bostoniensis 

14. Dorsal tentacles smooth or slightly wrinkled, not annulated 15 

15. Body ovate in outline, broad and somewhat depressed; with 

numerous cerata (up to 400 on each side) Aeolidia papillosa 

15. Body linear, narrow; with fewer cerata (not over 100 on each 

side) 16 

16. Body pale gray with white margins, and with 3 longitudinal 
reddish to russet interrupted stripes on head and anterior 

part of body (fig. 23) Cratena pi lata 

16. Body transparent white (internal organs may be yellowish or 
reddish), and without colored stripes on head and anterior 

part of body 17 

17. Anterior lateral extensions of the foot about half as long as 
width of foot; tips of cerata transparent with an opaque white 

ring just back of tip; anterior cluster of cerata not as dis- 
tinct and sharply set off as in the following; central core of 
cerata scarlet to rose red, varying in some specimens to choco- 
late (figs. 9-11) Coryphella rufibranchialis 

17. Anterior lateral extensions of the foot about as long as the 
width of foot; tips of cerata with patch of opaque white; the 
anterior cluster of cerata distinctly set off from second clus- 
ter forming a "ruff about the shoulders"; central core of cerata 
carmine red; cerata somewhat longer than in above species (figs. 

12-14) Coryphella pellucida 

18. Anterior lateral corners of foot bluntly angled (figs. 18, 19) 
Cuthona concinna 

18. Anterior lateral angles of foot rounded 19 

19. Cerata very numerous and crowded, irregularly set; lateral edge 
of each ceras produced into a wide, strongly undulating mem- 
brane (figs. 26, 27) Fiona pinnata 

19. Cerata not so crowded; cerata set in regularly defined rows; 

cerata circular in cross section and without lateral membrane 20 

20. Cerata fairly numerous (at least 30 to 50 on a side) 21 

20. Cerata few (4 to 10 or so on a side) 22 


Plate 20 


Figures 1, 2, 6-8, 18, 24-27 on Plates 20-22 
were drawn from Kodachrome transparencies taken 
by the author. Grateful acknowledgement is made 
to Miss M. Patricia Morse for inking most of the 
drawings. Figure 23 is redrawn from Verrill, 
the others from Alder and Hancock's Monograph. 
Figures of species that are satisfactorily il- 
lustrated in Miner (1950) have not been includ- 
ed in these plates. 

Figure 1. Pleurobranchaea tarda . Dorsal view of specimen from Sargas- 
sum , Vineyard Sound. 

2. P_. tarda . Ventro-lateral view of same. Ctenidium visible on right 

3. Acanthodoris pilosa . Dorso-lateral view. 

4. A. pilosa . Branchial plume. 

5. A. pilosa . Portion of mantle edge, highly magnified, showing the 
conical papillae. 

6. Corarobella (?) sp. Dorsal view of specimen from Sargassum , Vine- 
yard Sound. 

7. Corambella (?) sp. Ventral view of same, showing ctenidia-like 
gills at posterior end. 

8. Scyllaea pelagica . Lateral view of right side. Specimen from 
Sargassum, Vineyard Sound. 

All scale bars are 5 mm. 

Plate 20 


Plate 21 
Figure 9. Coryphella ruf ibranchialis . Dorso-lateral view. 

10. C. ruf ibranchialis . Ventral view of anterior end. 

11. c. ruf ibranchialis . Two cerata. 

12. Coryphella pellucida . Dorso-lateral view. 

13. C. pellucida . Ventral view of anterior end. 

14. C. pellucida . Two cerata. 

15. Eubranchus pallidus . Dorso-lateral view. 

16. Eubranchus exiguus . Dorsal view. 

17. Tergipes despectus . Dorsal view. Alder and Hancock noted that 
"The figures in this plate (Fam. 3, Plate 17) have inadvertently 
been reversed". The figure shown here is not reversed. 

18. Cuthona concinna . Dorsal view of specimen from Sakonnet Point, 
Rhode Island. 

19. C. concinna. Ventral view of anterior end. 

All scale bars are 5 mm. 

Plate 21 


Plate 22 
Figure 20. Cratena aurantia . Dorso-lateral view. 

21. C. aurantia . Ventral view of anterior end. 

22. C. aurantia . Two cerata. 

23. Cratena pi lata . Dorsal view. 

24. Facelina bostoniensis . Dorsal view of specimen from Tubularia , 
Woods Hole. 

25. F. bostoniensis . Ventral view of same specimen. 

26. Fiona pinnata . Dorso-lateral view of specimen from floating 
timber, Vineyard Sound. 

27. F. pinnata . Three cerata showing undulating membranous exten- 
sions of same. 

All scale bars are 5 mm. 

Plate 22 

25 •■ 

16i Shell-less Opiethobranchs 

21. Oral tentacles almost as long as dorsal tentacles; cerata 

long, cylindrical or slightly conical (figs. 20-22) .... Cratena aurantia 

21. Oral tentacles about half as long as dorsal tentacles; cer- 
ata ovoid ovate, much inflated, somewhat compressed (fig. 

15) Eubranchus pallidus 

22. Cerata 5 to 10 on a side, clavate, large and inflated, ta- 
pering abruptly to a point; body spotted with green or brown; 

tentacles banded with olive or brown (fig. 16) Eubranchus exiguus 

22. Cerata 4 (rarely 5) on a side, set in a single longitudinal 
row on each side; cerata smoothly (i.e. not abruptly) taper- 
ing; body transparent white, not spotted, sometimes striped 
anteriorly with two lateral reddish streaks; tentacles not 
banded (fig. 17) Tergipes despectus 


The list presented here includes most of the shallow water species of shell- 
less opisthobranchs known from the Woods Hole region, and is based both upon pub- 
lished records of Gould (1870), Verrill (1873), and Johnston (1934), and the col- 
lections of the author. Since we lack a monographic treatment of the group in 
this area, and since figures are helpful in identification, page references are 
made to Miner (1950) in the cases of species not figured in this manual. The syn- 
onymies given are not complete (some deletions in the interests of brevity have 
gone beyond the author's wishes, and for this the editor accepts responsibility). 
References to Alder and Hancock (1845-55) are given as A & H; Gould and Binney 
(1870) as G & B; Verrill and Smith (1873) as V & S. Additional synonymy is given 
by Johnston (1915), and Iredale and O'Donoghue (1923). 


Subclass Opisthobranchia 

Order Sacoglossa 

Elysia catula (Gould, 1870) . As Placobranchus catulus in G & B; as Elysiella 

catulus in V & S; Miner, p. 674. 
Elysia chlorotica Gould, 1870. Miner, p. 674. Found in salt and brackish water 

marshes . 

Order Notaspidea 

Pleurobranchaea tarda Verrill, 1880. See Verrill, 1882, Trans. Conn. Acad., 5_: 546. 
Dredged, 60-400 meters; also on floating Sargassum in Vineyard Sound. 

Order Nudibranchia 

Suborder Doridacea 

Acanthodoris pilosa (Abildgaard, 1789) . As Doris pilosa in A & H; as Doris bifida 
in V & S. Color various, white to yellow, brown, or even black. Under rocks, 
midtidal to laminarian zone; abundant sporadically in spring, also taken in sum- 
mer and fall; throughout New England. 

Corambella (?) sp„ Animals figured in figures 6 and 7 have been collected at Woods 
Hole on pilings and from Sargassum in Vineyard Sound. 

On chidoris a spera (Alder and Hancock, 1842) . As Doris aspera in A & H; as Doris 

pallida in G & B; as Onchidoris pallida in V & S; as Onchidorus aspera in Miner, 
p. 671. The spelling Onchidorus used by Miner and certain other authors is of 
incorrect gender, and has been emended by Winckworth (1932, J. Conchol. _19: 234, 
251). Under rocks, low intertidal to 60 meters. Probably feeds on encrusting 
bryozoans. All seasons; Bay of Fundy to Rhode Island. 

Shell-less Opisthobranchs 163 

Onchidoris fusca (Muller, 1776) . As Doris b i lame 1 lata in A & H and G & B; as Lamel - 
lidoris bilamellata in Miner, p. 672. Midtidal to 8 meters, on barnacle covered 
rocks; feeds on barnacles; often gregarious, breeding and laying eggs; all sea- 
sons, breeding in spring and fall. Bay of Fundy to Rhode Island. 

Palio lessonii (D'Orbigny, 1837) . As Polycera lessonii in A & H and V & S; Miner, p. 
673. Under stones, low intertidal; New Brunswick to Connecticut. 

Suborder Dendronotacea 

Dendronotus frondosus (Ascanius, 1774) . As D. arborescens in A & H, G & B, and V & 
S; Miner, p. 671. Often associated with hydroids, especially Tubularia . Large 
specimens (up to 8 cm) have been collected by the author from pilings at the 
east end of the Cape Cod Canal; specimens of 2-3 cm from Bay of Fundy to Long 
Island Sound af'all seasons. 

Idulia coronata (Gmelin, 1791) . As Dctocoronata in A & H, G & B, V & S, and Miner, 
p. 673. Associated with various hydroids, including Sertularia ; all seasons; 
Bay of Fundy to Long Island Sound. 

Scyllaea pelagica Linnaeus, 1761. On floating Sarqassum in Vineyard Sound. 

Suborder Eolidacea 

Aeolidia papillosa (Linnaeus, 1761) . As Eolis papillosa in A & H; as Aeolis papil - 
losa in G & B, and Miner, p. 670. This largest of New England eolids is some- 
times called the "common nudibranch", but it is sporadic in occurence. Found 
at all seasons, usually associated with sea anemones. Bay of Fundy to Woods 

Coryphella pellucida (Alder and Hancock, 1847) . As Eolis pellucida in A & H. Maine 
and Woods Hole; winter and spring. Not previously reported from New England. 

Coryphella ruf ibranchialis (Johnston, 1832) . As Eolis ruf ibranchialis in A & H; as 
Aeolis ruf ibranchialis in G & B. Balch's (1909) C. r. chocolata is but a color 
variety, intergrading with specimens with red cerata in the same area. The 
most abundant eolid of New England; among hydroids on rocks, intertidal to 200 
meters; winter, spring, and early summer; Bay of Fundy to Massachusetts. 

Cratena aurantia (Alder and Hancock, 1842) . As Eolis aurantiaca in A & H; as Mon - 

taqua gouldii in V & S. Associated with Tubularia . Abundant, summer and spring; 
New Hampshire to Woods Hole. 

Cratena pi lata (Gould, 1870) . As Aeolis pi lata in G & B; as Montaqua pilata in V & 
S. Abundant on Pennaria at Woods Hole in July and August. This is the species 
used by Kepner (1943, J. Morph. , 73_: 297-311) in his study of the manipulation 
of nematocysts by nudibranchs . 

Cuthona concinna (Alder and Hancock, 1843) . As Eolis concinna in A & H. A British 
species not previously reported from America; the author has taken it in Mass. 
and Rhode Island; Dr. Henry Russell has reported it from Cape Cod. 

Embletonia fuscata Gould, 1870. With red algae; New Hampshire to Woods Hole. 

Eubranchus exiquus (Alder and Hancock, 1848) . As Eolis exigua in A & H; as Aeolis 
despecta (in part) in G & B. Gould apparently confused Terqipes despectus and 
Aeolis exigua (see Verrill, 1882, Trans. Conn. Acad., 5_: 553, under Galvinja exi - 
gua) . On occasion E_. exiquus may lose its oral tentacles in the laboratory, and 
may then be confused with Embletonia fuscata . Associated with hydroids, Bay of 
Fundy to Woods Hole. Has been found breeding in MBL live cars in May. 

Eubranchus pallidus (Alder and Hancock, 1842) . As Eolis picta in A & H and Aeolis 
picta in G & B. Associated with hydroids, winter and spring; Bay of Fundy to 
Rhode Island. 

Facelina bostoniensis Couthouy, 1838. As Aeolis bostoniensis in G & B. Associated 
with Tubularia ; New Hampshire and Woods Hole region. 

Fiona pinnata Eschscholz, 1831. As Fiona nobilis in A & H; also F. marina of au- 
thors, and F. atlantica Bergh. Taken in August from Sarqassum and floating logs 
in Vineyard Sound. 

164 Shell-less Opisthobranchs 

Tergipes despectus (Johnston, 1835) . As Aeolis despecta (in part) in G & B; as Eo - 
lis despecta in A & H. Associated with hydroids, spring and summer; Bay of 
Fundy to Rhode Island. 


Alder, J. and A. Hancock, 1845-55. Monograph of the British Nudibranchiate Mollusca. 
Ray Society, London. 

Balch, F. N. , 1909. Notes on New England Nudibranchs II: A Spring Collecting Trip. 
Nautilus, 23.: 33-55. 

Eliot, C, 1910. The British Nudibranchiate Mollusca. Part VIII. (Supplementary 
to Alder and Hancock) . Ray Society, London. 

Gould, A. A. and W. C. Binney, 1870. Report on the Invertebrata of Massachusetts, 
2nd ed., vi-524 pp. Boston. 

Iredale, T. and C. H. O'Donoghue, 1923. List of the British Nudibranchiate Mol- 
lusca. Proc. Malacol. Soc . London, 1_5: 195-223. A useful checklist for synonymy. 

Johnston, C. W. , 1915. Fauna of New England, Part 13. List of the Mollusca. Occ. 
Papers, Boston Soc. Nat. Hist., pp. 1-231. 

Johnston, C. W. , 1934. List of Marine Mollusca of the Atlantic Coast from Labora- 
dor to Texas. Proc. Boston Soc. Nat. Hist., 40: 1-204. 

Miner, R. W. , 1950. Field Book of Seashore Life. xv-888 pp. Putnam, N. Y. 

Ohdner, N. H. , 1934. The Nudibranchiata. British Antarctic (Terra Nova) Exped. 
1910. Zool., 7_: 229-310. 

Ohdner, N. H. , 1936. Nudibranchia Dendronotacea. A Revision of the System. "Mel- 
anges Paul Pelseneer". Mem. Mus . Roy. d'Hist. Nat. Belg., Ser . 2, 3_: 1057-1128. 

Ohdner, N. H. , 1939. Opisthobranchiate Mollusca from the western and northern 

coasts of Norway. Det Kgl. Norske. Videnskabers Selskabs Skrifter, Nr . 1: 1-92. 

Thiele, J., 1929-31, 34, 35. Handbuch der systematischen Weichtierkunde. 2 vols. 
Gustav Fischer, Jena. 

Verrill, A. E. and S. I. Smith, 1873. Report upon the Invertebrate Animals of Vine- 
yard Sound, etc. Rept. U. S. Comm. Fish, for 1871-72, Part 1: 295-778, pi. 
1-38. Washington. 

Chapter XVI 2.65 


By Mary D. Rogick 

Entoprocta and Ectoprocta are zoologically distinct but because they bear super- 
ficial resemblance to each other they have been treated together by systematists, as 
"moss animals", Polyzoa, or Bryozoa ( sensu lato) . 

Entoprocts are minute, less than one centimeter tall. They are usually sessile, 
pseudocoelomate, and generally of soft texture. Their anus is within the ring of 
tentacles, which number 8 to 36, depending on the species. Tentacle number and stalk 
flexibility are important, so should be noted in living specimens wherever possible. 
The "heads" or calyces of some nod constantly. Some species are solitary, others 
colonial. Entoprocts may be found attached to rocks, algae, shells, or growing on 
various organisms as hydroids, sponges, Crustacea, ectoprocts and worms. 

KEY TO LOCAL ENTOPROCTA (partly after Osburn) 
(Figure references are to Plate 23 ) 

1. Individuals solitary, not colonial or stolonate, provided 
with a contractile stalk and enlarged basal disc (fig. 1) ; 
characteristically found on bodies of tube dwelling annelid 

or sipunculid worms Loxosoma spp. 

(Consult Prenant and Bobin for species differences) . 

1. Individuals colonial, arising from creeping stolons, forming 

low, whitish, simple traceries or sometimes denser tufts 2 

2. Stalks of individuals with distinct muscular dilations at 
bases, near junction with stolons, and sometimes elsewhere 

along stalk; stalks spineless Barentsia 3 

2. Stalks without such muscular dilations, tapering, usually 

with small spines on stalk and/or calyx (fig. 2) Pedicellina cernua 

3. Stalk thin walled and muscular, its distal end very flex- 
ible, actively curling into a loose spiral (fig. 3) Barentsia laxa 

3. Stalk heavier walled, straight and rigid 4 

4. Stalk wall appears as if punctured by irregularly spaced 

minute, cone shaped pores. Colony delicate, small Barentsia discreta 

4. Stalk wall without pores Barentsia major 

Family Loxosomatidae 

Loxosoma davenporti Nickerson, 1901. Reported (Nickerson) associated with the poly- 
chaete " Clymene producta " . It is not clear what maldanid is meant, possibly 
Maldane . 

Loxosoma minuta Osburn, 1912. May possibly be a Loxosome 11a ? Reported on Phasco - 
lion strombi (Sipunculoidea) . 

Family Pedicellinidae 

Barentsia discreta (Busk, 1886) . Inconspicuous but of world wide distribution. Found 

on algae, stones, hydroids, ectoprocts. See Rogick (1956). 
Barentsia laxa Kirkpatrick, 1890. Forms furry patches to half an inch high on Mer - 

cenaria shells which had been bored by the sponge Cliona. See Rogick (1948) . 

166 Entoprocts 

Barentsia major Hincks, 1888. Found on pilings, stones, shells, and around leg 

bases of spider crabs and Limulus . 
Pedicellina cernua (Pallas, 1771). On pilings and elsewhere, among bryozoans and 
" other creeping growths. 


Atkins, D. , 1932. The Loxosomatidae of the Plymouth area, including L. obesum , sp. 

nov. Quart. J. Micr. Sci., 75_: 321-391. 
Nickerson, W. S., 1901. On Loxosoma davenporti sp. nov. an endoproct from the New 

England coast. J. Morph., 17: 351-380, pi. 33. 
Osburn, R. C, 1912. The Bryozoa of the Woods Hole Region. Bull. U. S. Bur. Fish. 

(for 1910), 30: 203-266, pi. 18-31. 
Osburn, R. C, 1944. A survey of the Bryozoa of Chesapeake Bay. Chesapeake Biol. 

Lab. Publ. no. 63_: 1-59. 
Prenant, M. et G. Bobin, 1956. Bryozaires, Premiere Partie. Entoproctes, Phylac- 

tolemes, Ctenostomes. Faune de France, 60_: 1-398. 
Rogick, M. , 1948. Studies on marine Bryozoa. II. Barentsia laxa Kirkpatrick 1890. 

Biol. Bull., 94: 128-142. 
Rogick, M., 1956. Bryozoa of the U. S. Navy's 1947-1948 Antarctic Expedition, I-IV. 

Proc. U. S. Nat. Mus., 105: 221-317. 

Chapter XVII 167 


By Mary D. Rogick 

Ectoprocta, the "true" Bryozoa or "true" Polyzoa are a large and diverse group, 
composed of animals always individually minute, but organized into colonies of var- 
ied form. Since ectoprocts are important as fossils, much of the basic systematic 
work on the group has been done by paleontologists (see works of Canu and Bassler, 
particularly) ; consequently, reliance in taxonomy has been largely upon the exter- 
nal skeletal characters. 

In the field, the beginner may confuse ectoprocts with hydroids, sponges, sea- 
weeds, or compound ascidians, because of their growth forms. The colony texture var- 
ies from gelatinous, to membranous, to chitinous, to calcareous, depending upon the 
species . 

Identification to genus and species is often a matter of considerable difficul- 
ty. Original descriptions of some of the commonest species are too incomplete, too 
general and do not always include really distinguishing key characters. An origin- 
al description was sometimes so broad that several species could be included under 
one specific name and one species could fit with equal justification in more than 
one genus. Some genera, like Cellepora , Lepralia , Membranipora , and Smittina ac- 
tually became catch-alls for many diverse and hard-to-identify species. 

If possible, bryozoans should be studied alive, with their tentacles extended. 
Tentacle number is important in distinguishing some species. Use a compound micro- 
scope with lOOx to 280x magnification and direct (reflected) lighting for the study 
of calcareous or opaque species and transmitted lighting for transparent forms. Cal- 
careous colonies are sometimes calcined or treated with Eau de Javelle to remove 
organic matter from the skeleton. Calcining produces beautiful results but is ris- 
ky and may destroy the specimen, so is recommended only where ample material is 


"The study of ectoprocts is burdened by a large and fantastic terminology, much 
of it dating from a period when the structure of the animals was not understood. 
Hence the terminology lacks relation to terms employed for other groups of animals. 
Frequently the ectoproctologists seem to get themselves entangled in their own term- 
inology, using the same word (e.g., aperture) in several different senses". This 
statement by Dr. Libbie Hyman should be borne in mind when looking up bryozoan lit- 
erature. The general zoologist will find an illuminating discussion of ectoproct 
stru :ture and terminology in Vol. 5 of Hyman ' s "The Invertebrates". 

Ancestrula : Primary zoid produced by the metamorphosis of a larva. The rest of the 

colony develops from it by budding. 
Alveolus : Small cavity, pit, or fenced in area between zooecia, chiefly in Cyclo- 

Aperture : Variously used for any opening, sometimes for the zooecial orifice. Best 

to avoid its use. 
Areolae or areolar pores : One or more rows of pores around the periphery or margin 
of the zooecial front, often larger than other pores. Remainder of frontal 
wall, the central part, is usually imperforate if areolae are present. 
Autozoid: Normal, "typical", or sometimes asexual individual in a colony, if colony 
has male and female zoids (as in Hippothoa) . 

168 Ectoprocts 

Avicularium : Highly modified zoid, with or without a polypide but with muscles 

which operate its operculum, called the mandible. It may be adventitious , i.e. 
attached to some part of the parental zoid's front, or it may be vicarious , i.e. 
an independent unit, replacing a regular zoid in the colony, or be placed be- 
tween other zoids in the colony. Many sizes and shapes: larger than autozoids, 
smaller than autozoids; shaped like a bird's head, like the sole of a shoe, or 
round, elliptical, spatulate, triangular. May be imbedded in the calcareous 
wall of parent zooecium or swing freely on a stalk which is sometimes much long- 
er than the whole parent zoid. May even occur on ovicells of some species. 
Function unknown, although sometimes avicularia ward off other organisms or may 
keep other larvae from settling on the colony. 

Brood chamber : Used especially for cyclostomes. It is an enlarged space, compart- 
ment or ovicell in which zygotes develop into larvae. 

Brown body : A ball of brownish-orange tissue representing the remains of a degen- 
erating or degenerated polypide. Found either in empty or in inhabited zooecia. 

Cardelle : Denticle placed at each side within the orifice for hingement of the oper- 
culum; also called condyle . May be peg-like, unicusped, bicusped or even ledge- 

Communication pore : Opening in zooecial wall between adjacent zoids, for soft tis- 
sue connecting the two zoids . 

Connate : Firmly united or in close order, as in rows of zoids of Amathia. 

Cryptocyst : Shelf-like calcareous lamina extending inward from the front edges of 
the side walls, beneath the frontal membrane (like inward turned edges of pie 
crust) . In Anasca, especially in the membraniporid group. 

Dietellae: Blister-like chambers present in the basal-lateral walls of some Cheilo- 
stomes. They contain communication pores and are also called pore chambers. 

Distal : Direction of growth away from the ancestrula. That part of the zoid or 
colony that is farthest away from the ancestrula. 

Frontal side : The free or "ventral" side of an attached encrusting ectoproct, or 

the wall bearing the orifice and other decorations (avicularia, ovicells, pores, 
sculpturings, etc.) of a zoid. 

Gonozoid : Identifiable male or female zoid of a colony. Sometimes differ in size 
or in orifice from autozoids. Cyclostomes may develop brood chambers for 
developing young. Cheilostomes develop ovicells of various types. 

Gymnocyst : Peripheral calcified portion of frontal membrane in some Cheilostomes, 
developed especially in proximal region, not covered by membranous ectocyst. 
See figure 48 (Plate 26) . 

Heterozoid : Incomplete or highly modified zoid as opposed to autozoid. Includes 
avicularia, dwarf zoids, vibracula, gonozoids, some types of spines. Prefer- 
able to use term heterozooecium, since the zoid or soft parts may be vestigial. 

Lophophore : Circular or semicircular fold of tissue which bears the tentacles. 

Lyrula : A tooth, commonly anvil-shaped, low, of varying width, placed in the cen- 
ter of the proximal edge of the primary orifice. Characteristic of the "smit- 
tinid" orifice. Sometimes it may become very worn down. 

Mandible : The operculum closing the beak of an avicularium. 

Multiporous : Refers to a sieve-like plate in the lateral or distal walls of cal- 
careous Cheilostomes which serves as an interzoidal connection. 

Ooecium : Brood chamber containing the developing zygote or larva in Cheilostomes. 
Same as ovicell. 

Operculum : Membranous, chitinous or very rarely calcareous flap closing the ori- 
fice of Cheilostomata. Often works like a drawbridge, after tentacles have been 
withdrawn into the zooecium. 

Opesia : A large uncalcified membrane covered area on the front side of many Anas- 
can Cheilostomes (Membraniporids) . It surrounds the orifice and extends prox- 
imally. It is bordered by a cryptocyst. 

Oral avicularium : An avicularium that is either in the wall of the orifice or next 
to the orifice. Suboral would be just below or proximal to the orifice. Later- 
al oral would be to the side of the orifice. 

Ectoprocts lt>9 

Orifice : The zooecial opening through which tentacles are extruded from the zooe- 
cium. In Cheilostomes it is covered by an operculum. In some heavily calci- 
fied forms the original orifice (called the primary orifice ) may be hidden by 
a growing calcareous collar ( peristome ) whose free rim now forms a secondary 
orifice ( peristomice ) which may have an entirely different shape or appearance 
than that of the primary orifice. 

Orif icial collar : See Peristome and Orifice . 

Ovicell : General term for any structure serving to contain bryozoan larvae dur- 
ing their development, according to Bassler. May be of varied shapes and sizes 
and located conspicuously or inconspicuously on or within zooecia. Often placed 
at distal end of zoid, like a cap or hood. Sometimes placed in lateral posi- 

Peristome : An extension of the calcareous rim of the orifice in some Cheilostomes. 
See also Orifice . 

Pleurocyst : The calcareous frontal wall of some Cheilostomes. It is generally 
granulated and usually not porous over its central area but may have areolar 
pores around the edge. 

Polypide: The protrusible part of the bryozoan individual and that part which is 
suspended in the body cavity, namely: the tentacles, gut and associated mus- 
culature. Early workers on bryozoa thought that the bryozoan individual con- 
sisted of a box (zooecium) containing a soft individual (the polypide) . 

Pore chambers : See Dietellae . 

Primary orifice : See Orifice . 

Proximal : Direction toward origin of growth, or region nearest the ancestrula. 
That part of the zoid which is nearest its point of origin. 

Rhizoid: Same as radicle, a rootlike structure formed by zooecia for attachment of 

the colony to the substratum or to various objects. May be calcareous or chit- 
inous . 

Scutum : Large flabellate spine or shield in front of opesia; attached laterally. 

Secondary orifice : See Orifice . 

Septula : Communication pores between neighboring zoids. The pores may be single 

(uniporous) or grouped together in a sieve-like plate (multiporous) and located 
in the lateral and distal walls of zoids. 

Sinus : A slit or excavation or notch in the proximal part of the orifice. Espec- 
ially characteristic of the Schizoporellid group but may occur in other Cheilo- 
stomes . 

Stolon : Tubular strand, usually horizontal or recumbent (upright in Amathia ) from 
which either new zoids or other structures (peduncles or secondary stolons) may 
be budded. More common in Ctenostomata than in other orders, but sometimes 
present in the latter. 

Tremocyst : Perforated calcareous frontal wall of some Ascophoran Cheilostomes. If 
the wall is more or less perforated all over it is a tremocyst . If wall is per- 
forated only around the margin and imperforate in center, then it is a pleuro - 
cyst . 

Vibraculum : Highly modified chitinous or calcareous heterozoid found in Cheilo- 
stomes, resembling an avicularium whose mandible has been replaced by a very 
long bristle or chitinous whip, which may be considerably longer that the zo- 
oecium. The vibracular chamber contains powerful muscles for moving the whip. 

Zoarium : The entire bryozoan colony, composed of individuals called zoids or bryo- 
zoids or bryozoites . 

Zooecium : The external skeleton and body wall of the zoid. Originally coined for 
the compartments in which the polypides are housed. Term especially useful to 
paleontologists to whom only the hard skeletal structures are available for 

Zoid : A single individual of the colony. Term includes the zooecium and its con- 
tained polypide. 

170 Ectoprocts 


Preservation of soft or small calcareous specimens . Ryland (1962) suggests pre- 
servation of Ctenostomes in 4% sea-water formalin and preservation of calcareous 
forms in 70% alcohol. If calcareous forms are on large stones or shells, two courses 
are open to the worker: (1), to store the stones and intact colonies or (2), to burn 
off (calcine) the colonies and mount them on slides. In either case wash the stone 
or shell in fresh water and allow them to dry. Store dry. 

Calcining . To prepare permanent slide mounts of calcareous Ectoproct skeletons, 
one often resorts to calcining. To calcine (burn off the organic, membranous tis- 
sues), use an alcohol lamp or Bunsen burner and a geologist's blowpipe. Select a 
dry colony on a suitable rock or shell; blow with the blowpipe so as to direct a 
narrow jet of flame against the specimen. The colony will first blacken, then turn 
red hot and finally turn white. Protective glasses should be worn to protect the 
eyes and face from flying fragments of rock, shell or bryozoan. Caution: Do not 
burn the colony to a crumbly whiteness, but continue calcining only to the point 
where some white begins to show and the fine diagnostic pattern is retained. Col- 
ony fragments will lift up right off the rock and can be transferred on to the sur- 
face of very thick balsam on a slide. No coverslip is needed. If possible, mount 
a fragment of an uncalcined colony right alongside the calcined fragment for com- 
parison or in case the calcined fragment disintegrates. For further details, con- 
sult Rogick, 1945. 

Some people prefer to use a,bleach like Clorox (sodium hypochlorite) or Javelle 
water for cleaning and whitening specimens. 

(Figure references are to Plates 23-26) 

1. Zooecia are white calcareous tubes pitted with pores; orifice 
terminal, unconstricted in autozoids, i.e. not narrower than 
the autozoid, but small and constricted in the greatly swollen 
ovicells or brood chambers; tube tips free but rest of tube 
is either partly free or else immersed in the common zoarial 
crust; vibracula, avicularia and opercula absent; colonies may 
be arborescent, encrusting, or raised into stiff flattened lobes 
. . Order CYCLOSTOMATA ( =STENOLAEMATA Borg, 1926; =STENOSTOMATA Marcus, 1938) 2 

1. Zooecia (and gonozoids , if present) otherwise 4 

2. Erect, twig-like colonies articulated (having chitinous joints), 
attached at their origin to a primary calcareous disc; rhizoids 
present; zooecia slender; inflated vase-like ovicells densely 

pitted with pores (fig. 4) Crisia eburnea 

2. Colonies compact, discoid, wart-like, encrusting; the tubular 
zooecia open on the free surface out of the common crust; zo- 
oecial tubes arranged in series or fascicles radiating from 

a free central area; between them are adventitious tubes or 

incomplete partitions (aveoli or cancelli) ; brood chambers 

spread over and among several zoids 3 

3. Brood chamber roof overgrown with secondary alveoli, looking 

reticulated Lichenopora 

3. Brood chamber roof not covered thus, not reticulated (fig. 5) ... Disporella 

Ectoprocts 171 

Zooecia soft, gelatinous, membranous, corneous or leathery but not 
calcareous; zoids may be distinctly isolated or else closely pack- 
ed, encrusting, erect, stolonate, or some may burrow into mollusc 
shells; zooecial orifice terminal, closed by a puckering of the 
invaginated tentacle sheath usually, or occasionally by special 
structures (two lips in the Flustrellidae, a setigerous membrane 
in others (fig. 10), or an operculum in boring Penetrantiidae) ; 

true ovicells lacking but gonozoids may occur Order CTENOSTOMATA 5 

Colonies usually calcareous, but in some families are corneous 
or membranous; great variety of growth forms: encrusting, free- 
ly lamellate, arborescent, nodulate, stolonate, or reticulate; 
zooecia are rounded or angular chambers budding distally and/or 
laterally to form contiguous rows; zooecial orifice terminal or 
subterminal, more commonly at the distal end of zoid's frontal 
surface, and closed by a hinged operculum; avicularia and ovi- 
cells present in many species Order CHEILOSTOMATA 13 

Stolons or stolon-like extensions of zoids absent. Zoids are 
squat, opaque, their lateral walls fused together. Colony a 

gelatinous to leathery crust; common on coarse brown algae 6 

Stolons or stolon-like zoidal extensions present; zoids are 
membranous or chitinous, separate, tubular, more or less trans- 
parent and never leathery 7 

Colony rubbery, brownish. Closed orifice is a transverse 
bilabiate slit, purse-like (fig. 6); chitinous spines mount- 
ed on small pads (kenozooecia) that appear at edges of front- 
al wall, oftenest about the orifice Flustrellidra 

Colony a gelatinous to rubbery gray or brown crust sometimes 

arising into sac-like lobes; orifice at tip of a mound-like 

papilla closed by a puckering of the body wall (fig. 7) ; 

spines absent Alcyonidium 

Stolon false, non-septate, representing only the drawn out, 
narrowed, adherent proximal part of the zoid. Only transverse 
septum present is usually near point of origin of stolon. Re- 
mainder of zoid erect, tubular, with orifice squared 8 

Stolons genuine, divided into one or more segments by trans- 
verse septa. Erect zoids bud directly from stolon or from an 
intermediate peduncle 9 

Stolons long, zooecia tall (up to 3.8 mm); tentacles number 

8 to 22; gut very long (figs. 12, 13) Nolella 

Stolons shorter, zooecia very tall (up to 4.85 mm) and close 

together; new zoids may sprout from the lateral wall of erect 

zoids; 8 tentacles; funiculus long. In brackish water Victorella 

Stolon broad, tubular, often branches dichotomously; gizzard 

present 10 

Stolon slender, generally not dichotomous ; short or small lat- 
eral segments (peduncles) arise from it and give rise to zo- 
oecia 11 

172 Ectoprocts 

10. Zooecia very regularly disposed in a parallel series (forming 
a double row of closely packed, touching, parallel zoids) 
either in a continuous or an interrupted spiral around the 
stolon, or in palisade-like groups (fig. 11); colony arbor- 
escent Amathia 

10. Zooecia occur in irregularly clumped groups, or singly, along 
stolon. Zoids soft, flexible, rather transparent. When re- 
tracted their tips are squared, with corners reinforced. Mem- 
branous or setigerous collar shorter than in Aeverrillia .... Bowerbankia 

(1) Bowerbankia gracilis , the more common species, has 8 
tentacles (fig. 8) 

(2) Bowerbankia imbricata, less common, has 10 tentacles 
(fig. 10) 

11. Gizzard present. Zoids yellowish, horny, husk shaped, with 
membranous frontal area; occur in pairs near end of each 

internode (fig. 9) Aeverrillia 

11. Gizzard absent 12 

12. Zooecia clavate, with long slender stalk which attaches to 

the stolon peduncles; 12 to 21 tentacles Triticella 

12. Zooecia ovoid to cylindrical, originating from lateral 
branching sprouts of the stolon. Zooecia sometimes bunched 

up at nodes; 8 tentacles Valkeria 

13. Colony erect 14 

13. Colony recumbent or encrusting 21 

14. Avicularia, vibracula and true ovicells completely absent 17 

14. One or more of the above (avicularia, or vibracula or true 

ovicells) may be present 15 

15. Ovicells and avicularia may be present but both vibracula and 

scuta are absent 18 

15. Either vibracula or scuta, or both, may occur 16 

16. Vibraculum lacking but scutum present Tricellaria 

16. Vibraculum and scutum usually both present (fig. 40) Scrupocellaria 

17. Zoids tiny, single, erect, isolated, glassy, firm walled, and 
connected basally by stolonate extensions. Proximal part re- 
cumbent, swollen and punctate near where the upright spoon- 
shaped part diverges from it. Upright stalk reinforced by 
fine closely wound spiral thread. Colonies diffuse, white, 
inconspicuous but very common on bases of Bugula , hydroids and 

other colonial growths (fig. 14) Aetea 

17. Zoids biserial, back to back (fig. 44). New zoids and branches 
bud from the sides of the zooecia near distal end. Colonies 
yellowish, bushy. Opesia occupies about half the zooecial 

front and slants obliquely Eucratea 

18. Zooecia uniserial, budding from distal end and also from front- 
al wall just below the opesia; ovicell on dwarfed zoid; avicul- 
aria absent (figs. 15, 19) Scruparia 

18. Bird's head type avicularia usually present, zooecia biserial 

or multiserial rather than uniserial 19 

Ectoprocts 173 

19. Colony biserial; zooecia trumpet shaped, divided into three 
parts; opesia rounded, nearly terminal, at enlarged oblique- 
ly truncated end of zooecium. Upward facing orifice surround- 
ed by 4 to 8 extremely long spines. Avicularia and ovicells 
lateral (fig. 16) Bicellariella 

19. Zooecia not trumpet shaped but more tubular. Opesia very 
large, occupying from half to nearly the whole of the zooecial 

frontal surface 20 

20. Base of zooecium transverse at its place of origin dorsally 

and proximally; zooecia multiserial (fig. 52) Dendrobeania 

20. Base of zooecium strongly forked at its place of origin dor- 
sally and proximally. Zoarium biserial to narrowly multiser- 
ial (figs. 17, 18) Bugula 

Two species are common at Woods Hole: 

(1) Bugula simplex (fig. 17), formerly called B. f labellata , 
tan in color with flattened, somewhat fan-like fronds. 
Common in such protected places as the Eel Pond. 

(2) Bugula turrita (fig. 18), yellow to orange-brown; colony 
of conical form with a marked spiral or whorled arrange- 
ment of branches. Found in more exposed situations. 

21. Colony a fragile calcareous lace; zooecial front has a large 
uncalcified membranous area (opesia) ; side walls and their 

inturning ledges (cryptocyst) calcified 23 

21. Zooecial frontal wall more extensively calcified 22 

22. Frontal wall covered by two rows of calcareous flattened ribs 
(costae) , more or less fused, with rows of pores or perfora- 
tions where ribs did not quite meet Cribrilin a 

(1) Cribrilina annulata (fig. 28): Frontal costae and rows of 
pores regular and distinct; avicularia absent; ooecia 

(2) Cribrilina punctata (fig. 29): Ooecia large; avicularia 
present at sides of orifice. 

22. Frontal wall not costate but well calcified except for the 

orifice and possible pores 26 

23. Ovicells absent; avicularia wanting in most species 24 

23. Ovicells and avicularia present; avicularia located proxi- 
mally or laterally on front wall; opesial spines often 

present 25 

24. Avicularia absent; spines usually present around opesial 
border (figs. 21, 22, 23). Well developed gymnocyst 

usually present Electra 

24. Avicularia absent from most species. No calcareous spines 
on opesial walls but tubercular processes may occur at zo- 
oecial corners. Cryptocyst may develop into a proximal 
shelf under frontal membrane but a regular gymnocyst (cal- 
careous outer front wall) is wanting or greatly reduced 
(fig. 20) Membranipora group 

The genus Membranipora is a temporary" dumping genus" for 
hard-to-identify "open-faced" species. Species are shifted 
from it and its daughter genera Conopeum , Acanthodesia , etc. 
and then returned to the mother genus. Membraniporan classi- 
fication is still very fluid. 

174 Ectoprocts 

25. Several large blister-like pore chambers present in basal- 
lateral walls of zoid; they are punctured by communication 
pores (fig. 24) Callopora 

25. Pore chambers absent; instead, zooecial walls contain sev- 
eral uniporous or multiporous septules (pore plates) Teqella 

26. Primary orifice without a proximal median tooth, the lyrula 27 

26. Primary orifice rounded to subcircular, with median proximal 

lyrula; cardelles (denticles) usually present 35 

27. Orifice semicircular, with straight proximal border, without 
a cardelle (denticle) in each corner; median suboral asco- 
pore present; frontal wall with many pores; avicularia and/ 

or ovicells on some zoids (fig. 25) Micropore 11a 

27. Orifice otherwise; special ascopore absent 28 

28. Proximal border of orifice forms a shallow cradle-like si- 
nus as wide or wider than the rest of the orifice. Front- 
al wall coarsely perforated all over 29 

28. Proximal border of orifice forms a sinus narrower than the 
rest of the orifice (a "keyhole" orifice) ; frontal wall ei- 
ther areolate or with pores all over 30 

29. Orifice somewhat bell shaped because of sinus width (fig. 30); 

avicularia and ovicells absent Cryptosula 

29. Orifice rounded; perforate ovicells and occasional avicular- 
ia present Hippodiplosia 

30. Orifice with beaded distal vestibular arch separated from 
the wide proximal sinus by a broad bicusped or bifid car- 
delle at each side. Ovicell with large uncalcified front- 
al area 

Hippoporina contracta (fig. 26) but not other Hippoporinae 

30. Orifice arch not beaded; ovicells and cardelles not as 

above 31 

31. Avicularia absent; frontal wall non-porous; ovicells with 
small number of pores; ovicelled zoids of smaller size 

than autozoids 32 

31. Avicularia present; frontal wall variously porous 33 

32. Male zoid with rounded notched orifice like that of auto- 
zoid, but smaller; cardelles unicusped or bicusped; female 
gonozoid orifice differently shaped from that of autozoid. 
Female gonozoid on same face of colony as autozoids. Col- 
ony small, vitreous, uniserial to multiserial (fig. 27) Hippo thoa 

32. Female gonozoid on back surface of autozoids. Colony uni- 
serial or biserial Haplota 

33. Frontal wall a tremocyst, i.e., perforated all over by pores; 

cardelles small or wanting; ovicells generally with pores 34 

33. Frontal wall imperforate except for areolar pores; ovicell 
evenly perforated by pores; small oral avicularium on asym- 
metrical suboral umbo; other avicularia elsewhere, of var- 
ious shapes and sizes (fig. 36) 
Schismopora and some of the other Celleporae 

Ectoprocts 175 

34. Avicularium present in midline proximal to orifice Schizomavella 

34. Avicularium not in midline below orifice but located 

elsewhere about orifice or frontal (figs. 32-35) Schizoporella 

35. Avicularia present 36 

35. Avicularia absent; frontal wall imperforate except for 

areolar pores; ovicell imperforate Mucronella 

36. A median suboral avicularium present 37 

36. Avicularium not median nor suboral; frontal wall areolate 
but not perforated centrally; avicularia may be peripheral 

(fig. 31) Parasmittina 

37. Suboral avicularium median and longitudinally directed 38 

37. Suboral avicularium transverse or obliguely placed on a 
suboral umbo just in front of or partly obscuring the ori- 
fice Rhamphos tome 11a 

38. Frontal wall and ovicell perforated by pores all over the 
central area; sometimes the median suboral avicularium be- 
comes incorporated into the peristomial collar in old zo- 

oecia Smittina 

38. Frontal wall imperforate except for areolar pores 39 

39. Ovicell center area perforate Smittoidea 

39. Ovicell usually imperforate, or at most with an occasional 

pore Porella 

(Figures of genera mentioned in key are not mentioned here) 


Order Cyclostomata (= Stenolaemata or Stenostomata) 

Crisia cribraria Stimpson, 1853. Rare. At Crab Ledge. 

Crisia denticulata (Lamarck, 1836) . Doubtful identification. 

Crisia eburnea (Linnaeus, 1758). Delicate white brittle upright slender branching 

sprigs. Common on algae, especially Chondrus crispus , drif tweed and holdfasts. 

Disporella hispida (Fleming, 1828) . Flat rounded white calcareous patches resemb- 
ling lichens; about 1/8 inch in diameter; edges crinkly, center part with 
slightly raised tubes or jagged projections. On algae, hydroids, bryozoa and 
stones . 

Lichenopora verrucaria (Fabricius, 1780). On stems of Bugula . Eucratea , hydroids, 
and on shells and stones. About 1/8 inch in diameter. 

Oncousoecia diastoporides (Norman, 1868). Fig. 37; Stomatopora of some authors. 
Rare, from Crab Ledge. 

Tubulipora atlantica (Johnston, 1847). At Crab Ledge, on stones and shells. 

Tubulipora flabellaris (Fabricius, 1780). Fig. 43. Uncommon, on shells and stones 
at Crab Ledge and near Nantucket. 

Tubulipora liliacea (Pallas, 1766). Uncommon, on algae, eel grass, shells and stems 
of hydroids and Bugula , in Vineyard Sound. 

Order Ctenostomata Busk, 1852 

Aeverrillia armata (Verrill, 1873) . On piles and seaweed ( Laminaria and Phyllo - 
phora ) . 

176 Ectoprocts 

Aeverrillia setigera (Hincks, 1887) . On hydroids and such algae as Chondrus and 
Ascophyllum . Indistinguishable in field from Aeverrillia armata . 

Alcyonidium gelatinosum (Linnaeus, 1767). Questionable. 

Alcyonidium hirsutum (Fleming, 1828) . Vineyard Sound, on algae. 

Alcyonidium parasiticum (Fleming, 1828). Vineyard Sound, Crab Ledge and No Man's 

Alcyonidium polvoum (Hassall, 1841). A. mytili of Osburn ' s 1912 paper. Encrusts 
piles, barnacles, stones, algae, shells, Libinia crab, and even skate egg cas- 
es. Color very variable, from gray to yellow to red to brown. 

Alcyonidium verrilli Osburn, 1912. Rare at Vineyard Sound. 

Amathia vidovici (Heller, 1867). A. dichotoma of Osburn ' s 1912 paper. Common, on 
piles, rocks, oyster shells and algae. 

Anquinella palmata Van Beneden, 1845. Rare, mud encrusted. Not in key. 

Bowerbankia gracilis var . caudata (Hincks, 1877). On stones, shells, ascidians, 
and on stems of hydroids, bryozoa and algae. 

Bowerbankia gracilis Leidy, 1855. On piles, stones, and about 18 species of sea- 
weeds . 

Bowerbankia imbricata (Adams, 1800) . The least common Bowerbankia , indistinguish- 
able in field from B. gracilis . Colonies pinkish in breeding season (July 
and August) because of reddish larvae. Found on algae ( Chondrus , Fucus, As - 
cophyllum , Corallina ) . 

Flustrellidra hispida (Fabricius, 1780) . Once known as Flustra or Flustrella . 

Forms a brown, rubbery crust on such algae as Ascophyllum , Chondrus , Fucus , 

Nolella blakei Rogick, 1949. On Perophora from Lagoon Pond, Martha's Vineyard. 

Triticella elongata (Osburn, 1912). Commensal on legs, shells or branchial cham- 
bers of such crabs as Callinectes sapidus , Libinia , and on pinnotherid crabs 
in Chaetopterus tubes. 

Triticella pedicellata (Alder, 1857) . Recorded as Vesicularia f amiliaris in Os- 
burn 's 1912 paper. On algae. 

Valkeria uva (Linnaeus 1758) . From Vineyard Sound, on hydroids and bryozoa. 

Victorella pavida K ent, 1870. Membranous, soft, white tracery or tuft; in brack- 
ish water. 

Order Cheilostomata Busk, 1852 
Suborder Anasca Levinsen, 1909 

Aetea anguina (Linnaeus, 1758). Tiny but common on stems of algae, and animals, 

and on stones and shells. 
Aetea recta Hincks, 1861. The Aetea sica of Rogick and Croasdale's 1949 paper. 

On about 18 algal species. Pinkish larvae found in July and August. 
Amphiblestrum f lemingii (Busk, 1854). Fig. 49; Membranipora f lemingii in Osburn, 

1912. On shells, stones and algae. Not in key. 
Bicellariella ciliata (Linnaeus, 1758). On piles, stones, shells and hydroids. 

Embryos in ovicells in July and August. 
Bugula avicularia (Linnaeus, 1758). Not in key; see fig. 50. 
Bugula cucullif era Osburn, 1912. Not in key; see Rogick and Croasdale (1949). 

On algae ( Fucus , Laminaria , Rhodymenia and Phyllophora ) along with Aetea and 

Crisia . 
Bugula simplex Hincks, 1886. This has been extensively used experimentally under 

the name of B. f labellata . Forms thick yellow-orange tufts in protected 

places such as floats in Eel Pond and piles elsewhere. 
Bugula turrita (Desor, 1848). Occurs in more exposed situations than B. simplex , 

and easily recognized by its spiral growth. Also much used in laboratory 

studies. Larvae released from late June throughout August. Grows on at 

least 16 algal species. 

Ectoprocts 177 

Bugulopsis peachii var ■ berinqia Kluge, 1952. Fig. 47. Cellularia peachii in 
Osburn 1912. Rare; on shells and on Dendrobeania murrayana . Not in key. 

Caberea ellisii (Fleming, 1818). Fig. 41; on shells and pebbles. Not in key. 

Callopora aurita (Hincks, 1877) . Formerly in Membranipora . Small white colonies 
on rocks and less commonly on such algae as Phyllophora , Phycodrys an< 3 hold- 
fasts of Laminaria . 

Callopora craticula (Alder, 1857). {- Membranipora formerly). On shells and 
stones . 

Callopora lineata (Linnaeus, 1767). (= Membranipora in Osburn 1912). Rare; on 
shells, stones and algae. 

Cauloramphus cymbaeformis (Hincks, 1877) . Fig 38; formerly in Membranipora . En- 
crusting stalks of hydroids and Dendrobeania murrayana . 

Cellaria fistulosa (Linnaeus, 1758). Not in key. 

Conopeum reticulum (Linnaeus, 1767). Fig. 39; Membranipora lacroixii of Osburn, 
1912, and Rogick and Croasdale, 1949. Delicate encrusting lace on rocks, 
shells and less frequently on such algae as Ascophyllum , Fucus and Phyllo - 
phora ; sometimes covers an area of several square inches. 

Cribri lina annulata (Fabricius, 1780). Rare; on stones, shells, and algae ( Phyco - 
drys and Laminaria ) . 

Cribri lina punctata (Hassall, 1841) . Not common, but has been found encrusting 
shells, pebbles and 7 algal species. 

Dendrobeania murrayana (Johnston, 1847). Height 0.5 to 1.5 inches; common in 
outer waters from dredged shells and pebbles. 

Electra crustulenta (Pallas, 1766) . As Membranipora monostachys in fig. 29b of 
Plate XXII, Osburn (1912). 

Electra hastinqsae Marcus, 1938. As Membranipora monostachys in fig. 29a of Plate 
XXII, Osburn (1912). Mostly on rocks and shells but occasionally on Fucus , 
Laminaria , and even on gill chambers of the spider crab Libinia . 

Electra pilosa (Linnaeus, 1767). Very common on Laminaria ; occurs also on about 

16 other algal species, as a fine calcareous lace, one layer thick, sometimes 
a foot in length. 

Eucratea loricata (Linnaeus, 1758) . Bushy phytoid colonies up to 10 inches high 
in outer waters (Crab Ledge, Nantucket, and No Man's Land). Formerly called 
Gemellaria . 

Membranipora tenuis Desor, 1848. Cryptocyst forms a jagged shelf that covers the 
proximal half of the opesia. Encrusts stones and shells. 

Membranipora tuberculata (Bosc, 1802) . (= Membranipora tehuelca ) . Exceedingly 
abundant on Sargassum , sometimes on Laminaria and drif tweeds. 

Scruparia ambigua (d'Orbigny, 1841) . Fig. 19. Found on Bugula turrita and about 
11 algal species ( Laminaria , Fucus , etc.). 

Scruparia chelata (Linnaeus, 1758). Fig. 15. Not common, but has been reported 
on bryozoa, hydroids and algae, and also on piles. 

Scrupocellaria scabra (Van Beneden, 1848). Rare; on shells, stones and in drift. 

Tegella arctica (d'Orbigny, 1851). (= Membranipora) ■ Colonies one inch in dia- 
meter, on shells and stones. 

Tegella armifera (Hincks, 1880) . Membranipora arctica var. armifera of Osburn 
(1912) . 

Tegella unicornis (Fleming, 1828). (= Membranipora ) . Encrusts dredged shells. 

■rncellaria gracilis (Smitt, 1867). Menipea ternata in Osburn's 19l2 paper. At- 
taches to shells, stones, hydroids and bryozoans . 

Suborder Ascophora Levinsen, 1909 

Cellepora avicularis Hincks, 1860. Cellepora americana in Osburn, 1912. On algae 
and stems of hydroids and bryozoans. 

Cellepora canaliculata Busk, 1881. On hydroids and bryozoans. 

Cellepora dichotoma Hincks, 1862. On such algae as Chondrus , Gracilaria and Phyllo - 
phora . See fig. 36. 

178 Ectoprocts 

Cryptosula pallasiana (Moll, 1803) . (= Lepralia ) . Living colonies an orange color, 
especially around the periphery. Colonies flat, calcareous, about 2 cm in dia- 
meter; common on rocks and shells; also occur on about 11 species of algae such 
as Laminaria , Fucus , Ascophyllum , Ulva , etc. 

Cylindroporella tubulosa (Norman, 1868). Fig. 51. Porina tubulosa in Osburn, 1912. 
On stones and shells, in outer waters; not common. Not in key. 

Haplota clavata (Hincks, 1857). Fig. 42. (- Scruparia ) . Rare; on Dendrobeania mur - 
rayana and Eucratea loricata . 

Hippodiplosia americana (Verrill, 1875). (- Lepralia ) . On shells and stones; col- 
onies white to reddish. 

Hippodiplosia pertusa (Esper, 1796). (Formerly Lepralia ) . White to reddish cal- 
careous colonies of considerable extent on rocks and shells. 

Hippoporina contracta (Waters, 1899). Lepralia serrata of Osburn. White to buff- 
colored calcareous colonies encrusting rocks, shells and some algae (Phyllo - 
phora ) . Zoids small and crowded. The beaded orifice is distinctive. 

Hippothoa divaricata Lamouroux, 1821. On stones, shells and occasional algae; rare. 

Hippothoa hyalina (Linnaeus, 1767) . Exceedingly common and cosmopolitan species, 

encrusting red and brown algae especially but also found on stones, shells, hy- 
droids and bryozoans . Forms tiny glistening white to irridescent patches about 
2 or 3 mm in diameter, usually twining around small algal stems or in protect- 
ed spots, as on holdfasts. Embryos plentiful in July and August. 

Micropore 11a ciliata (Pallas, 1766) . On rocks, shells and 5 algal species. 

Microporella ciliata var . Stella ta (Verrill, 1875). On shells. 

Mucronella immersa (Fleming, 1828) . Mucronella peachii in Osburn, 1912. On stones 
and shells; occasionally on algae. 

Mucronella ventricosa (Hassall, 1842). Rare; on stones and shells. 

Parasmittina nitida (Verrill, 1875) . Smittina trispinosa var. nitida of Osburn. 

Colonies very fine grained; form lightweight multilayered porous nodules sever- 
al inches in diameter; color gray to sulfur yellow. Abundant in dredgings. 

Parasmittina trispinosa (Johnston, 1838). On stones, shells, and occasionally on 
algae. Colonies whitish to yellow. 

Porella acutirostris Smitt, 1867. On shells and stones; colonies rounded, pattern 
often of great regularity. At Crab Ledge. 

Porella concinna (Busk, 1852). Common at Crab Ledge on stones and shells. 

Porella proboscidea Hincks, 1888. White or yellow frilly bilaminate colonies ris- 
ing erect from a base that encrusts shells, stones, and the ascidian Boltenia , 
sometimes several inches high; from Nantucket Shoals, No Man's Land, and Crab 

Porella propinqua (Smitt, 1867). On shells and hydroid stems. 

Rhamphos tome 11a bilaminata (Hincks, 1877). On hydroid stems. 

Rhamphos tome 11a costata Lorenz, 1886. Colony encrusts stems of various kinds, ris- 
ing frill-like to a height of one-half inch; at Crab Ledge and Nantucket Shoals. 

Rhamphos tome 11a ovata (Smitt, 1868). Rare, encrusting stones and shells. 

Schizomavella auriculata (Hassall, 1842) . (= Schizoporella ) . Colorless to yellow 
to reddish colonies encrust stones, shells and occasionally hydroid stems; at 
Crab Ledge and Nantucket Shoals. 

Schizoporella biaperta (Michelin, 1841-42) . (= Stephanosella) . Multilaminate ruf- 
fly reddish-orange colonies encrust piles, stones, shells, hydroid stems and 
9 algal species ( Chondrus , Fucus , Laminaria , etc.). 

Schizoporella unicornis (Johnston, 1847) . Multilaminate red calcareous colonies 

encrust shells, stones, piles, worm tubes and 6 algal species ( Chondrus , Fucus , 
Laminaria , etc) . 

Smittina majuscula Nordgaard, 1905. Smittia porifera of Osburn ' s 1912 paper. Col- 
ony encrusts stones, shells and stems of various kinds; off Nantucket and Crab 

Stomachetosella sinuosa (Busk, 1860). Fig. 45. (= Schizoporella ) . Circular red, 

purple or brown colonies encrust stones and shells at Crab Ledge. Not in key. 

Ectoprocts 179 

Umbonula arctica (Sars, 1851). Fig. 46. Mucronella pavonella of Osburn ' s 1912 
paper. Colony encrusts stones and shells or forms fan shaped expansions on 
stems of hydroids, etc. Not common. Not in key. 


Bassler, R. S., 1953. Bryozoa: 253 pp. Part G of R. C. Moore ' s "Treatise on Inver- 
tebrate Paleontology." Geological Soc . of America, univ. of Kansas Press. Law- 
rence, Kans . Exceedingly useful for both recent and fossil genera. 

Borg, F., 1944. The Stenolaematous Bryozoa. Further Zool. Results Swedish Antarc- 
tic Exped. 1901-1903. Ill (5): 1-276, 16 plates. Includes many comments and 
revisions on other than Antarctic species. 

Harmer, S. F., 1923. On Cellularine and other Polyzoa. Jour. Linn. Soc. Zool. 

35 : 293-361; 3 plates. Very important morphological paper settling some taxo- 
nomic questions. 

Harmer, S. F., 1957. The Polyzoa of the Siboga Expedition, Part IV, Cheilostomata 
Ascophora. . .Resultats Siboga-Expedit. XXVIIId (Livr. 145): 641-1147; 33 plates. 

Hincks,T, 1880. A history of British Marine Polyzoa, 2 vols., London, England. A 
classic reference. 

Hyman, L. H., 1959. "The Invertebrates", vol V., Smaller Coelomate Groups. McGraw- 
Hill Book Co., New York, N.Y. Ectoprocta on pp. 275-515. The most comprehen- 
sive compilation on the subject. 

Kluge, G. A., 1962. Mshanki Severnih Morei SSSR. (Bryozoa of the Northern Seas of 
the USSR). (In Russian). Fauna USSR #76, Akademia Nauk USSR, Zool. Institute, 
Leningrad. 584 pp. Very good figures. 

Lagaaij, R. , 1952. The Pliocene Bryozoa of the Low Countries. Mededelingen van de 
Geologische Stichting, Ser. C, V(5) : 1-233; 26 plates. 

Marcus, E-, 1940. Mosdyr (Bryozoa eller Polyzoa). Danmark ' s Fauna, Dansk Natur- 
historisk Forening 46, 401 pp.; G. E. C. Gads Forlag, Kizfoenhavn, Danmark. Ex- 
cellent handbook for identification. 

Osburn, R. C, 1912. The Bryozoa of the Woods Hole Region. Bull. U. S. Bur. Fish. 
(1910), 30: 205-266; pi. 18-31. 

Osburn, R.C., 1950 .Bryozoa of the Pacific Coast of America. Part 1. Cheilostomata- 

Anasca. Allan Hancock Pacific Exped. 14(1): 1-269, pi. 1-29. Univ. of Southern 
Calif. Press, Los Angeles, Calif. 

Osburn, R. C, 1952. Part 2. Cheilostomata-Ascophora Ibid . , 14(2): 271-611; pi. 

Osburn, R. C, 1953. Part 3. Cyclostomata, Ctenostomata, Entoprocta, and Addenda. 
Ibid . , 14(3): 613-841; pi. 65-82. 

Prenant, M. et G. Bobin, 1956. Bryozoaires, Iere Partie, Entoproctes, Phylactolames, 
Ctenostomes. Faune de France, 6_0: 1-398. Very usable. 

Rogick, M., 1940. An ecological effect of the New England hurricane. Ohio Jour. 
Sci., 40 (3) : 163-167. 

Rogick, M., 1945. Calcining specimens. Amer . Biol. Teacher, £5: 66-70. 

Rogick, M. , 1949. Studies on marine Bryozoa, IV. Nolella blakei , n. sp. Biol. Bull., 
97 : 158-168. 

Rogick, M., 1956. Bryozoa of the U. S. Navy's 1947-48 Antarctic Expedition, I-IV, 
Proc. U. S. Nat. Mus., Smithsonian Inst., 105 (3358) : 221-317; 35 pi. 

Rogick, M. and H. Croasdale, 1949. Studies on marine Bryozoa, III. Woods Hole re- 
gion Bryozoa associated with algae. Biol. Bull., 96: 32-69. 

Ryland, J. S., 1960. The British Species of Bugula (Polyzoa). Proc. Zool. Soc. Lon- 
don, 134 (1) : 65-105. Exceedingly useful for identification of some troublesome 
Bugulae . 

Ryland, J. S., 1962. Biology and identification of intertidal Polyzoa. Field Stud- 
ies, I_(4) : 19 pp. 

Silen, L., 1944. On the formation of the interzoidal communications of the Bry- 
ozoa. Zooloogiska Bidrag fran Uppsala. Band 22^: 433-488. Excellent account of 
the various types of interzoidal communications, pores, chambers, etc. 

180 Ectoprocts 

Soule J. D., 1957. Two species of Bryozoa Ctenostomata from the Salton Sea. Bull. 

So. Calif. Acad. Sci . , 56(1): 21-30. Adult Nolella blakei and Victorella pavida . 
Zoological Record: Bryozoa (Polyzoa) Section. An annual publication dating from 

1864 to the present; indispensable for any worker in the group. 

Plate 23 


Figures 1, 7, and 11 after Osburn 1912; figs. 2, 
4-6, 8-10 after Rogick and Croasdale 1949; figs. 
3 and 12-13 after Rogick 1949; all redrawn by 
Mrs. Emily Reid 

Fig. 1. Loxosoma davenporti . 

2 . Pedicellina cernua . 

3. Barentsia laxa . 

4. Crisia eburnea : note inflated ovicell and joints or nodes. 

5. Disporella hispida , a complete small colony. 

6. Flustrellidra hispida , portion of colony; note spines and 
slit-like closure of orifices. 

7. Alcyonidium verrilli , portion of colony; note puckered clo- 
sure of orifices. 

8. Bowerbankia gracilis , zoids with retracted tentacles. 

9. Aeverrillia armata , portion of colony; note paired zooecia on 
short peduncles, and the 4 terminal spines on each zooecium. 

10. Single polypide of Bowerbankia imbricata with tentacles extend- 
ed; note setigerous collar directly below the tentacles, here 
constricting the tentacle sheath. 

11. Amathia vidovici ; note close-set spiral bands of zoids. 

12. Nolella blakei , retracted individual, very young, with squared 
orifice . 

13. Nolella blakei , very young zoid, with tentacles extended. The 
four basal extensions are "false" stolons. The bottom right 
represents the attenuated proximal end of the shown zoid. The 
other three are cut off by septa from the base of the shown 
zoid and likewise represent the proximal extensions of their 
own zoids. 

Plate 23 



Plate 24 


Figures 15, 16, 20 and 21 after Osburn (1912); 
rest after Rogick and Croasdale (1949), redrawn 
by Mrs. Emily Reid 

Figure 14A, B. Aetea recta , note proximal recumbent parts of zoids, 
connected by stolonate extensions. 

15. Scruparia chelata; a zoid at left bears ooecium. Opesia oblique 
and shorter than in §_. ambiqua . 

16. Bicellariella ciliata : A, portion of colony; B, ovicell borne 
on a zoid; C, avicularium with serrated beak. 

17. Bugula simplex : small part of branch, with 3 bird head avi- 
cularia and 3 overhanging ovicells. 

18. Bugula turrita : small part of branch, shows 4 ovicells and one 
bird head avicularium. Note spines on zooecia. 

19. Scruparia ambiqua , portion of colony arising from basal attached 
row of zooecia. Opesia parallel to back wall and longer than in 
S^. chelata . 

20. Membranipora tuberculata : note large opesia and pairs of tubercles. 

21. Electra crustulenta , a more or less spineless species. 

22. Electra pilosa , a lightly calcified spiny species. Note porous 
gymnocyst (frontal proximal wall) , and spines bordering opesia. 

23. Electra hastingsae , a species with delicate spines, sometimes 
lacking or broken off. 

24. Callopora aurita, part of a colony showing 9 zooecia, 10 ovicells 
(with triangular front area) and 13 avicularia (some smaller than 
others) . 

25. Microporella ciliata: note the spine bordered hemispherical ori- 
fice under which is a small median crescent shaped ascopore. 
Four globose ovicells at bottom conceal orifices. 

26. Hippoporina contracta: note bifid denticles and beaded arches of 
the orifices. Ovicells are not pictured but note spatulate avi- 
cularia on the two extreme right zoids, and areolar pores. 

Plate 24 


Plate 25 


Figure 30 after Osburn, rest after Rogick 
and Croasdale, all redrawn by Mrs. Emily Reid 

Fig. 27. Hippo thoa hyalina , one ooecium at upper right of group. 

28. Cribrilina annulata , no ooecia shown. 

29. Cribrilina punctata , three ooecia shown at right in group. 

30. Cryptosula pallasiana . 

31. Parasmittina trispinosa , portion of colony without ooecia. 
Note avicularia beside orifices. 

32. Schizoporella unicornis , portion of colony without ooecia. 
Note avicularia beside orifices. 

33. Schizoporella unicornis , zooecia with ovicells. 

34. Schizoporella biaperta , heavily calcified portion of colony; 
note avicularia close to orifices. 

35. Schizoporella biaperta , portion of less heavily calcified 
colony, without ovicells. 

36. Cellepora dichotoma , portion of colony, with ovicells. Note 
avicularium borne on umbo on front of each zooecium. 

Plate 25 




186 Plate 26 


Figures modified from indicated sources; 
redrawn by Mary Rogick. Some of the gen- 
era figured are not in the key. 

Fig. 37. Colony fragment of Oncousoecia diastoporides , after fig. 12A of Osburn, 1912. 

38. Five zoids and two pedicellate avicularia of Cauloramphus cymbaeformis . after 
figs. 36, 36A of Osburn, 1912. 

39. Five zoids of Conopeum reticulum , after fig. 9 of Rogick, 1940. The large ope- 
sia is here blacked in. 

40. Branch of Scrupocellaria scabra , after Plate 6, fig. 7 of Hincks, 1880, showing 
frontal and lateral avicularia, and scuta (aperture shields) . 

41. Caberea ellisi from dorsal aspect, showing three long vibracula, after PI. 8, 
fig. 7 of Hincks, 1880. 

42. Haplota clavata from dorsal aspect, showing ovicell arising from back of zoid 
on branch at left; from fig. 107 of Marcus, 1940. Detail of aperture (orifice) 
above at right . 

43. Very young colony of Tubulipora flabellaris . showing the "primitive disc" at 
point of colony origin; after PI. 64, fig. 2 of Hincks, 1880. 

44. Eucratea loricata , showing back-to-back zooecia, from fig. 16 of Osburn, 1912. 

45. Three zoids and an ovicell (at bottom) of Stomachetosella sinuosa . from fig. 
51 of Osburn, 1912. 

46. Umbonula arctica. zoid with 2 adventitious avicularia and the characteristically 
large orifice; from fig. 16 of Osburn, 1912. Note the areolar pores and small 
denticle or umbo (not a lyrula) . 

47. A single internode of Bugulopsis peachii var . beringa after fig. 20 bis of Os- 
burn, 1912. Note dark "joints" at bottom and top. 

48. A composite diagram, modified from fig. 116 of Bassler, 1953, showing two mem- 
braniporid zoids, and a vicarious (independent) avicularium at right. The low- 
est zoid shows a pointed adventitious frontal avicularium. This zoid's non- 
porous frontal surface is an olocystal gymnocyst. The upper zoid has a frontal 
wall with pores, a tremocystal gymnocyst. The cryptocyst is the shelf immediate- 
ly framinj the opesia. The cap shaped structure overhanging the distal end of 
the top zoid is an ovicell. 

49. An ovicelled zoid of Amphible strum f leminqii similar to Callopora except for 
the design on the ovicell; after fig. 38 of Osburn, 1912. 

50. Dorsal surface of a Bugula avicularia branch, showing the forked proximal ends 
of zoids (where zoids originate distally and dorsally) , after PI. 10, fig. 2 of 
Hincks, 1880. 

51 A, 51B, 51C. Cylindroporella tubulosa , after figs. 43, 43A, 43C of Osburn, 1912. 
Fig. 51A is a side view of an ovicelled zoid; fig. 51B is the frontal view of 
another ovicelled zoid; 53C shows several bottle shaped autozoids . 

52. Dorsal surface showing the transverse proximal ends of Dendrobeania murrayana ; 
after Pi. 14, fig. 6 of Hincks, 1880. 

Plate 26 

188 Chapter XVIII 


Keys axe provided for the common members of four classes of living echinoderms 
(the Crinoidea are not represented at Woods HoleX These classes may be recognized as 
such without difficulty, except in the case of the very worm-like synaptid holothur- 
ians. Keys revised with the help and advice of John M. Anderson. Figure references 
are to Plate 27. 

I. Class Asteroidea 
(Sea stars or starfishes) 

1. Arms bear prominent spines; tube feet in 4 rows Asterias 2 

1. Arms bear only numerous minute spines; tube feet in two rows 3 

2. Arms tend to be blunt and cylindrical; skeleton firm; jaws 

of major pedicellariae (on adambulacral spines) bx'oad and blunt 

(fig. 1); colors usually dark, brown to greenish black; common 

Asterias forbesi 

2. Arms pointed and somewhat flattened; skeleton rather soft; 

usually a line of spines on upper surface of each arm; jaws of 

major pedicellariae tapering and pointed (fig. 2); colors lighter, 

yellowish to lavender; rare south of Cape Asterias vulgaris 

3. Five arms; usually crimson above, yellow below; small; more com- 
mon north of Cape Henricia sanguinolenta 

3. Nine to eleven arms; up to 25 cm across; found north of the Cape 

Solaster endeca 

II. Class Ophiuroidea 
(Brittle stars, serpent stars) 

1. Lateral spines on arms short 2 

1. Lateral spines on arms prominent 3 

2. Disc granulated, with no distinct scales; 2 pairs of slits leading 

into genital bursae at base of each arm (fig. 3) . . . . Ophio derma brevispina 

2. Disc covered with small scales; only one pair of slits leading 
into genital bursae at base of each arm (fig. 4) ; found only 

north of the Cape Ophiura robusta 

3. Three lateral arm spines on each joint (fig. 5); small, gray and 

white Amphipholis squamata 

3. Five or six lateral arm spines on each joint (fig. 6); color 

variable, often bright ; arms often banded Ophiopholis aculeata 

III. Class Echinoidea 
(Sea urchins, sand dollars) 

1. Flat and disc-like; very numerous minute spines Echinarachnius parma 

1. Globular, with prominent spines 2 

2. Color purplish, brownish, or blackish, spines long and stout; 

the common sea urchin about Woods Hole Arbacia punctulata 

2. Color greenish or yellowish; spines short, slender, and more 

numerous; much more common north of the Cape 

Strongylocentrotus drobachiensis 

Echinodermata 189 

IV. Class Holothuroidea 
(Sea cucumbers) 

1. Body wall muscular and opaque; typical tube feet present; 10 

branching tentacles surround mouth 2 

1. Body wall thin, transparent, showing 5 longitudinal ("radial") 
muscle strands; 12 pinnately branched tentacles about mouth; 
tube feet absent; worm-like in appearance, tending to adhere 

to fingers when handled 4 

2. Tube feet scattered over body, not confined to five adambulacral 

rows Thyone briareus 

2. Tube feet mostly in 5 rows Cucumaria 3 

3. Large (up to 30 cm); color some shade of brown; north of Cape 

Cucumaria frondosa 

3. Small (up to 6 cm); color grayish or pale; south of Cape 

Cucumaria pulcherrima 

4. Color whitish; tentacular digits 4-6 pairs plus terminal digit 
(variable) ; usually in sand (see note in check list) 

Leptosvnapta tenuis 

4. Color pinkish; tentacular digits 2-4 pairs plus terminal digit 
(variable) ; usually in gravel or under stones (see note in 
check list) Leptosvnapta roseola 

Note: the two synaptid species can be distinguished by the following internal 

characteristic: In the calcareous ring of L. tenuis each radial plate contains 
a hole for the passage of the radial nerve (fig. 7) ; in L. r oseola these plates 
are not pierced but are notched anteriorly (fig. 8). 

I. Class Asteroidea 

Asterias austera Verrill, 1895. Found only offshore in deep water. Not in key. 

Asterias forbesi (Desor, 1848) . "Common starfish" south of the Cape. 

Asterias tenera Stimpson, 1862. Not in key. This "slender armed starfish" occurs 
in deeper water (6-40 meters) both north and south of the Cape, but is not re- 
corded from Vineyard Sound. 

Asterias vulgaris Verrill, 1866. "Northern starfish"; South of the Cape this spe- 
cies occurs only in deeper water. 

Henricia sanguinolenta (O. F. Muller, 1776). "Blood starfish", formerly known as 
Cribrella sanguinolenta . The large yolky eggs are brooded beneath the mother; 
development is direct, without a free larval stage. 

Solaster endeca (Retzius, 1783). Does not occur at Woods Hole; found north of the 

II. Class Ophiuroidea 

Amphipholis (formerly Amphiura ) squamata (Delle Chiaje, 1828). One of the commonest 
ophiuroids of the region, but small and inconspicuous; generally in gravel, 
stones, or shell. Development direct, in brood pouches of parent. 

Gorgonocephalus agassizi (Stimpson , 1853). Not in key. Remarkable for its dichoto- 
mously branching arms, the "basket star" or "spider" occurs in northern waters. 
Known to the tip of the Cape and Nantucket Shoals. It is occasionally brought 
into Woods Hole by fishermen. 

Ophioderma (formerly Ophiura ) brevispina (Say, 1825) . Very common in protected, 
grassy bays such as Lagoon Pond. 


Plate 27 


From specimens or redrawn from various 
sources, all by Bruce Shearer. 

Figure 1. Asterias f orbesi . outline of a major pedicellaria, 
showing blunt form (after Coe) . 

2. Asterias vulgaris . outline of major pedicellaria, showing 
more elongated form (after Coe) . 

3. Ophioderma brevispina . underside of junction of arm with 
disc, to show the two pairs of openings of the genital 

4. Ophiura robusta . underside of junction of arm with disc, 
showing one pair of openings of genital bursae (after 
Clark) . 

5. Amphipholis squamata , lateral view of a portion of an arm, 
to show the three lateral arm spines per joint. 

6. Ophiopholis aculeata , lateral view of a portion of an arm, 
showing six lateral arm spines per 'joint. 

7. Leptosvnapta tenuis , radial and interradial pieces of cal- 
careous ring, the radial pierced for the passage of the 
radial nerve (after Heding) . 

8. Leptosvnapta ( Epitomapta ) roseola , radial and interradial 
pieces of the calcareous ring, the radial notched anter- 
iorly (after Heding) . 

Plate 27 



Ophiopholis aculeata (Linnaeus, 1767). Uncommon; the "daisy brittle star"; may be found 

in exposed rocky areas, as at Cuttyhunk. 
Ophiura robusta (Ayres, 1851) . Found only north of the Cape. 

III. Class Echinoidea 

Arbacia punctulata (Lamarck, 1816). "Purple" or "common sea urchin". One of the best 

known sources of embryological material. 
Echinarachnius parma (Lamarck, 1816). "Sand dollar"; taken by dredging on sandy bottom. 
Strongylocentrotus drobachiensis (0. F. Miiller, 1776. "Green sea urchin"; taken rather 

rarely in Vineyard Sound, but is the common (and only) urchin north of the Cape. 

IV. Class Holothuroidea 

Caudina arenata (Gould, 1841) . Not in key. This apodous (tube-foot-less) holothurian 
much resembles a smooth skinned sipunculid with one end drawn out into a sort of 
tail. It is common in sand north of Boston and has been reported off Cuttyhunk. 

Cucumaria frondosa (Gunnerus, 1770) . This large sea cucumber may be expected only 
north of the Cape. 

Cucumaria pulcherrima (Ayres, 1854) . A small gray form; may occasionally be taken, 
or washed up in numbers, on Nobska and Stony Beaches after severe storms. 

Leptosynapta ( Epitomapta ) roseola (Verrill, 1873). Heding (1928, p. 323) considers 
this form sufficiently distinct to be given subgeneric or even generic rank as 
Epitomapta roseola . 

Leptosynapta inhaerens , see L eptosynapta tenuis . 

Leptosynapta tenuis (Ayres, 1851) . This species is almost always referred to at Woods 
Hole as L. inhaerens (0. F. Miiller, 1738). But on the basis of a careful compari- 
son, Heding (1928) concludes that the "L. inhaerens " of Woods Hole is not of the 
same species as the "L. inhaerens "of Europe and that the next available name, ten - 
uis , should be used. The fact of this difference should be borne in mind if ex- 
perimental results obtained on L. inhaerens in Europe and America are compared. 

Thyone briareus (Lesueur, 1824) . A well known experimental animal, collected in very 
soft mud in shallow bays. 

Thyone scabra Verrill, 1873. Rare, dredged. Not in key. 

Thyone unisemita (Stimpson, 1851) . Rare, dredged. Not in key. 


Clark, H. L, 1904 

for 1902, 22 
Coe, W. R., 1912. 


Heding, S 


The Echinoderms of the Woods Hole region. Bull. U. S. Fish. Com. 


Echinoderms of Connecticut. Conn. State Geol. Nat. Hist. Survey, 
19: 1-152. 

E., 1930. The holothurians of the western part of the Atlantic Ocean. 
Mus. Comp. Zool., Harvard, 7_1(3) : 43-266, pi. 1-24. 

G., 1928. Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16, 
Synaptidae. Vidensk. Medd. Dansk Naturhist. Foren., 8_5: 105-323. 

Chapter XIX 193 


The term "protochordates" today includes the Phylum Hemichordata, and two subphyla 
of the Chorda ta, namely: 

A. Subphylum Cephalochordata, with Amphioxus and its kin, not represented locally. 

B. Subphylum Urochordata (- Tunicata) which includes: 

1. Class Ascidiacea, the sessile tunicates, well represented in the Woods Hole 
fauna and much used in experimental work. 

2. Classes Thaliacea and Larvacea, which are pelagic or planktonic, and are not 
covered in these keys. 


The only common hemichordate of Woods Hole is the well known Saccoglossus kowal - 
ewskyi (A. Agassiz, 1873) . This occurs intertidally or in shallow water in fine muddy 
sand where its presence is marked by little piles of fine, stringlike castings. The 
wormlike body is exceedingly fragile and difficult to collect undamaged. It is easily 
recognized by the characteristic whitish proboscis, orange collar, and brownish body 
(plate 28, fig. 1). 

The development of S. kowalewskyi is direct, without drastic metamorphosis, from a 
ciliated larva. Some confusion has resulted from the claim of Agassiz that S_. kowal - 
ewskyi arose from the metamorphosis of a planktonic "tornaria" larva. The fact that 
tornaria larvae do appear in local waters indicates that some other species may exist 
in not too distant, but probably deeper, waters. 

Agassiz described our common species as Balanoqlossus kowalewskyi ; this was reas- 
signed to Dolichoglossus by Spengel in 1893, in his well known work on the Enteropneusta 
of the Gulf of Naples. However, Van der Horst, in his revision of 1939 (Bronn's "Tier- 
reich", IV:4:2:2) adopted the prior name of Saccoglossus (published by Schimkewitsch in 
1892), but long overlooked as a consequence of the paper being in Russian). Since there 
is no question as to the species kowalewskyi (or the variant spelling kowalevskii ) , ex- 
perimentalists should use the specific name and not merely the generic name in reporting 
work done on Saccoglossus kowalewskyi at Woods Hole, in order to avoid possible confu- 
sion to foreign readers. 



Class Ascidiacea 

This section has been compiled from material written and contributed by Drs. Donald 
J. Zinn and Donald P. Abbott, whose assistance is much appreciated. The key is intended 
only for identification of ascidians in the Woods Hole region. Information about the 
anatomy and ecology of these animals may be found in Berrill (1950) and further material 
on their taxonomy in Van Name (1945). Embryological details are discussed at some length 
by Berrill (1936). Figure references on ascidians are to plate 28. 

Ascidians show various degrees of colony formation: " simple " ascidians develop sing- 
ly from eggs and do not bud; individuals in this group are relatively large. Colonial 
ascidians can reproduce by budding, and form aggregations of several to many small indi- 
viduals. In " social " ascidians, such as Perophora , the individuals are distinct from 
each other and are connected only by stolons at their bases (figs. 2, 3). " Compound " 
ascidians, such as Botryllus , have minute individuals embedded in a common test -or tunic 
which often has a characteristic growth form (figs. 7, 8). 

194 Ascidiacea 

It is often very difficult to see the anatomical features of fresh, unrelaxed as- 
cidians, since they contract strongly when dissected. For purposes of laboratory study 
of anatomy, or if necessary for identification, relaxed and properly preserved speci- 
mens should be used. Place animals in pans of sea water to which is added Epsom salts 
(MgS04) , about one heaping teaspoon per liter. After 12-24 hours of anaesthetization, 
fix for at least an hour before study in 5-10% formalin made up with sea water. This 
treatment permits easy observation of features nearly impossible to discern in delicate, 
contractile living forms or in contracted preserved material. Specimens that are kept 
for more than a couple of weeks should be transferred through 2 changes of 70% alcohol. 
Large specimens should have slits made in them to facilitate penetration. Fresh speci- 
mens will usually be easier to study if hardened a little by previous transfer to strong 
alcohol for a few days. 

In using the following key, note that the branchial or inhalent siphon (morpho- 
logically the mouth) is considered anterior. Dorsal is defined by the position of the 
brain and subneural gland, which lie close to the surface between the siphons. The 
gonads in solitary ascidians are typically conspicuous structures, each consisting of 
a sac-like ovary on the surface of which may be located the lobes of the testes. 


1. Colonial ascidians (individuals are small and connected by 

stolons or embedded in common test) ; 

1. Simple ascidians (individuals usually of good size, and al- 
though they may occur in clusters are not organically con- 
nected to each other) 

2. Colony thin, flat, encrusting; zooids colorful, and arranged 
in stellate or subcircular clusters or "systems" (figs. 7, 8); 
zooids of each system with branchial apertures opening inde- 
pendently, atrial apertures directed medially and opening into 
a common cloaca 1 cavity in the tunic Botryllus schlosseri 

2. Colony not as above 3 

3. Colony consisting of numerous tiny (2-4 mm diameter) greenish 
individuals, connected to one another only by basal stolons 

(social ascidians) (figs. 2,3) Perophora viridis 

3. Colony not as above; individual zooids completely embedded in 

matrix of the common test (compound ascidians) 4 

4. Forming flat encrusting colonies 2-4 mm thick and up to more 
than 10 cm in diameter; test smooth on surface, opaque because 
of large numbers of minute stellate calcareous spicules which 
are white, yellowish, or reddish in color; no postabdomen pre- 
sent Didemnum candidum 

4. Form of colony otherwise; test not containing stellate calcar- 
eous spicules, though sometimes with encrusting or embedded 

sand grains; postabdomen present, containing the gonads 5 

5. Colony in the form of massive 'meaty" plates, lumps, or lobes up 
to several inches long and attached by one edge; tunic very 
firm; stomachs of zooids bearing about 12 longitudinal ridges 

(fig. 5) Amaroucium stellatum 

5. Colony not as above; stomachs of zooids bearing 20 or more long- 
itudinal ridges (fig. 4) 6 

6. Colonies low, globular, oval or flat topped, attached by a nar- 
row base Amaroucium constellatum 

6. Colonies consisting of globular masses built up of radially ar- 
ranged, transparent, finger-like lobes, which may contain encrusting 
and embedded sand Amaroucium pellucidum 



7. Branchial siphon with 8 lobes, atrial siphon with 6; body 

elongate, up to several inches long; test grayish, gelatinous, 

transparent enough when expanded to reveal the pharynx with 5-7 

longitudinal muscle bands on each side and the main loop of 

the gut, posterior to the pharynx, basally (fig. 6) Ciona intestinalis 

7. Branchial siphon with 4 or 6 lobes; body and test not as above 8 

8. Branchial aperture (also atrial aperture) square or 4-lobed; 
dissection shows pharyngeal stigmata straight, elongate, and 

not arranged in spiral or circular fields; no kidney present .... STYELIDAE 9 

8. Branchial aperture usually 6-lobed, atrial aperture usually 
4-lobed; dissection shows pharyngeal stigmata more or less 
curved and arranged in circular or spiral fields; kidney pre- 
sent on the right side MOLGULIDAE 10 

9. Test leathery, tough, irregular, yellowish or brownish in 
color; when collected often feels like a rough lump of gristly 
consistency; 2 gonads on each side of the body (figs. 9, 10); 

pharynx with 4 longitudinal folds on each side Styela partita 

9. Test thin, tough, membranous, pinkish to bright red in life, 
attached by a relatively large area basally; body depressed 
with long axis usually parallel to the substrate; 1 gonad on 
the right side (fig. 12), none on the left (fig. 11); pharynx 
with one longitudinal fold on the right side, none on the left 
Dendrodoa carnea 

10. Adults live in soft sand or mud substrates, unattached to 

larger objects 11 

10. Adults do not live free in soft sand or mud, but occur attached 
to larger objects such as gravel, rocks, floats, pilings, marine 

plants, shells 12 

11. Siphons arise close together; body covered with minute fib- 
rous processes which hold a layer of mud about the body; 
dissection shows only one gonad, this present on the left 
side (figs. 13, 14); pharynx lacking longitudinal folds but 
bearing 7 large internal longitudinal vessels on each side 
Bostrichobranchus pilularis 

11. Siphons separated at bases by a space greater than 1/5 of 
body circumference (figs. 15, 16); dissection shows 2 gonads, 
one on each side of the body; pharynx bearing longitudinal 
folds on each side, each fold marked by a small group of in- 
ternal longitudinal vessels Molgula arenata 

12. Very common, attached to eel grass, floats, pilings, etc., 
in bays; long axes of both gonads straight (figs. 17, 18) 
Molgula manhattensis 

12. Less common; long axes of one or both gonads conspicuously 

curved 13 

13. Gonads on both sides an inverted U-shape, leaving open ends 
of oviducts pointed away from the base of the atrial siphon 

(figs. 19, 20) Molgula complanata 

13. Left gonad with an S-shaped or W-shaped curve, right gonad 
with a straight axis; both gonads with open ends of oviducts 
curved to point toward the base of the atrial siphon (figs. 
21, 22) Molgula citrina 


Plate 28 


Figures 1, 3, and 9 from life; rest redrawn 
from Van Name (1945) ; all by Bruce Shearer. 

Fig. 1. Saccoglossus kowalewskii , whole animal, from life. 

2. Perophora viridis . individual, seen from left side. 

3. Same, colony from life, to show "social" growth habit. 

4. Amaroucium constellatum , zooid with larvae in atrial cavity. Note 
the numerous stomach ridges. 

5. Amaroucium stellatum , region of stomach, to show the fewer and 
simpler ridges. 

6. Ciona intestinalis , diagram of body seen from left, to show posi- 
tion of gut and gonads; branchial sac omitted. 

7. Botryllus schlosseri , single zooid seen from left side. 

8. Same, small colony from life, showing investment in common tunic. 

9. Stye la partita , diagram of body as seen from left and as if wall 
transparent, branchial sac omitted, showing gut and gonads. 

10. Same, seen from right side, showing gonads. 

11. Dendrodoa carnea, view of left side showing gut. 

12. Same, view of right side, showing gonad. 

13. Bostrichobranchus pilularis , diagram of body seen from left, showing 
gut and gonad. 

14. Same, from right, showing renal sac. 

15. Molgula arenata , body from left, showing gut and gonad. 

16. Same, body from right, showing renal sac and gonad. 

17. Molgula manhattensis , body from 18. Same, from right, 

19. Molgula complanata , body from left. 20. Same, from right. 

21. Molgula citrina , body from left. 22. Same, from right. 

Plate 28 

198 Ascidiacea 


Order Aplousobranchia 

Family Synoicidae 

Amarouciuni stellatum Verrill, 1871, Massachusetts to Florida. "Sea Pork"; viviparous. 
Amaroucium constellatum Verrill, 1871. New Hampshire to Florida. Good embryological 

material. Viviparous. 
Amaroucium pellucidum (Leidy, 1855) . Massachusetts to Florida. In 10-20 meters on 

sand, shells, or gravel in tidal currents. Viviparous. 

Family Didemnidae 

Didemnum candidum lutarium Van Name, 1910. Van Name (1945) considered the form pre- 
valent at Woods Hole, and extending from southern Maine to the south side of Cape 
Cod, to be a northern geographical race of D. candidum Savigny, 1816. This may 
prove erroneous (see Millar, 1962. Studies Fauna Curacao ... 8_: pp. 62-66), but 
the familiar name D. candidum is retained until further investigation shows the 
necessity of a change. 

Order Phlebobranchia 

Family Cionidae 

Ciona intestinalis (Linnaeus, 1767). One of larger ascidians of the United States, and 
one of the best known and most widely distributed of all ascidians. Greenland to 
Rhode Island. Oviparous. 

Family Perophoridae 

Perophora viridis Verrill, 1871. Although small, its transparent test makes it favor- 
able for observation of heartbeat, etc. Southern Cape Cod to southern Florida. 
"Sea grape"; viviparous. 

Order Stolidobranchia 

Family Botryllidae 

Botryllus schlosseri (Pallas, 1766). Very common; colonies most often found encrust- 
ing in variety of colors on hydroid stems and eel grass leaves. Portland, Maine 
to West Coast of Florida; viviparous. 

Family Styelidae 

Styela partita (Stimpson, 1852) . Formerly well known as Cynthia partita ; the eggs are 
classical embryological material. Common on wharf pilings, often with other ascid- 
ians, from Massachusetts Bay to West Coast of Florida. "Sea peach" (note that the 
name "sea peach" is also used for the long -stalked ascidian Boltenia, often cast up 
on beaches north of Cape Cod, and the large solitary Halocynthia aurantium of more 
northerly waters) . 

Dendrodoa ( Styelopsis ) carnea (Agassiz, 1850). Pinkish to bright red body spreads out 
on surfaces of stones, dead mussel shells, etc., to which animal is attached. New- 
foundland to Long Island Sound. Viviparous. 

Ascidiacea 199 

Family Molgulidae 

Bostrichobranchus pilularis (Verrill, 1871) . Size and transparency of test depend on 
environment. Found on or in mud or sand from one meter to greater depths. Tadpole 
larvae not formed. St. Lawrence River to Florida. 

Mogula arenata Stimpson, 1852. Commonly unattached; buried; regular, symmetrically 

circular in profile (except between siphons) , disk-like when viewed from above and 
generally covered with coat of sand grains. South Shore of Cape Cod to Cape May, 
New Jersey. Tadpole larvae not formed. Oviparous. 

Molgula citrina Alder and Hancock, 1848. Usually attached to hard objects or pilings; 
vary greatly in appearance and shape; are transluscent dull olive. Gulf of St. 
Lawrence to Narragansett Bay, Rhode Island. Viviparous. 

Molgula complanata Alder and Hancock, 1870. Unusually long synonymy (see Van Name) ; 

sand and shell encrusted and more or less globular body attached by slender fibrous 
processes to stones, shells or hard sand from low water to about 300 fathoms. Gulf 
of St. Lawrence to Martha's Vineyard. Viviparous. 

Molgula manhattensis (DeKay, 1843). Commonest ascidian of this area. Nearly globular 
body with test irregularly hirsute and usually papillated near siphons; attached 
to others of same species in large groups or to wharves, pilings, ship bottoms, eel- 
grass in shallow, polluted, brackish or ordinary sea water. Portland, Me. to N. E. 
Texas (southern Florida?) Oviparous with larvae developing outside of body of 

Molgula robusta (Van Name, 1912) . Not in key. Differs from M. manhattensis in a) 
lacking free-swimming larval stage, b) larger average size, c) living at 17-25 
meters on compacted mud-sand bottoms in areas of tidal currents, unattached and 
usually on left side, d) long pointed teeth on margin of posterior part of dorsal 
lamina, and e) reddish purple eggs. Woods Hole Harbor and Vineyard Sound. Ovipar- 
ous . 


Berrill, N. J. 1936. Studies in tunicate development. Part V. The evolution and class- 
ification of ascidians. Phil. Trans. Roy. Soc, London Ser. B., Vol. _226i 43. 

Berrill, N. J. 1950. The Tunicata. London, Ray Society, 354 pp. 

Grasse, P. P., Ed. 1948. Traite de Zoologie. Tome XI. Echinoderms, Stomocordes, Pro- 
corde's. pp. 895-919. 

Van Name, W. G. 1945. The North and South American Ascidians. Bull. Amer. Mus . Nat. 
Hist. 84: pp. 1-476, pi. 1-31. 



Among non-scientific or "common" names of invertebrates, a relatively small num- 
ber are truly vernacular, that is, firmly established in the speech of the region 
and generally known to natives. Some true vernacular names as "horseshoe crab" and 
"quohog" may actually be more stable than scientific names, and are quite specific. 
But other vernacular terms such as "rock crab" or "barnacle" lack precision because 
each applies to more than one species in the region, not distinguishable by the lay- 
man, or because even if only one species is indicated in a particular region, the 
same name is used for other species elsewhere. Many so-called common names are not 

vernacular, but may represent anglicized versions of scientific names thus we 

speak of "an ameba", "a hydra", "planar ians" , or "isopods"; such names are general- 
ly learned in school and are better known to the scientific than to the general pub- 
lic. They are useful, but not in referring to particular species. Then there are 
the sometimes barbarous renditions from scientific specific names, such as "Morton's 
egg cockle" for " Laevicardium mortoni" . Names of this sort are characteristic among 
groups of animals widely studied by amateurs. In the case of birds and fishes, such 
names have become fairly well standardized. Among invertebrates, such are most com- 
monly used for shell bearing molluscs, and since Abbott's "American Seashells" and 
similar works provide easy reference, we have included few names of this sort. 

The use of synthetic "common" names does nothing to aid the beginning student 
and is to be discouraged in scientific writing. Likewise, the use of non-specific 
vernacular names, unless accompanied by the proper scientific name, can lead to 
great confusion in the literature and should especially be avoided by comparative 
physiologists and ecologists. Far better to use an "old" but unmistakable scienti- 
fic name like Nassa obsoleta , Nereis limbata , or Mactra solidissima than a meaning- 
less vernacular term such as "the clam", "the rock crab", or "the sand worm". 

In the following glossary, we have attempted to include the common vernacular 
names characteristic of southern New England, as well as names which have been im- 
ported and have taken root among the Woods Hole scientific community. We make no 
claims of completeness, and would appreciate criticisms. For the great majority of 
invertebrates, there are no common names, vernacular or otherwise. 

"Common name " 

Scientific approximation 

" Common name " Scientific approximation 

Acorn barnacles . . . Balanus , Chthamalus 

Acorn worms Hemichordata 

Bamboo worms .... Clymenella , Maldanidae 
Barnacles . . Cirripedia (shelled types) 

Beach fleas, hoppers Amphipoda 

(various species, esp. 

of family Talitridae) 

Bearded sponges . . Microciona prolifera 

Big fiddler crabs Uca minax 

Black fiddler crabs Uca pugnax 

Black quohogs Arctica islandica 

Blood clams . . Anadara (various species) 
Blood stars . . . Henricia sanguinolenta 

Blood worms Glycera 

Blue crabs Callinectes sapidus 

Boring sponges . Cliona (several species) 

Bread sponges Halichondria 

Brittle stars Ophiuroidea 

Broken back shrimp Hippo lyte 

Bubble shells Haminoea, etc. 


fiddler crabs .... Uca pugilator 
Channeled whelks 

Busycon cana lieu latum 

Cherrystone clams. . young Mercenaria 

Chitons . Polyplacophora (Amphineura) 

Clams . . Various Bivalvia (Mollusca) 

not including the generally 

recognized mussels, scallops, 

oysters, etc. 

Clam worms Nereis 

Columbus's crabs . . . Planes minutus 
Comb jellies Ctenophora 

Corals Madreporaria and some 

other colonial Anthozoa 
Dead-man ' s fingers . Alcyonium carneum 

Dog whelks Thais lapillus 

Edible mussels .... Mytilus edulis 

Eel grass Zostera 

Feather -duster worms . . . Sabellidae 



Fiddler crabs Uca (3 species; 

Finger sponges Haliclona oculata 

Fish lice Branchiura ( Arqulus ) 

Flatworms Platyhelminthes 

Goose barnacles . Lepas (several species) 

Grass shrimp Hippo lyte 

Green crabs Carcinus maenas 

Green sea urchins 

. . . . Strongvlocentrotus drobachiensis 

Gribbles Limnoria 

Gulf weed Sarqassum 

Gulf weed crabs Planes minutus 

Hard-shelled clams Mercenaria 

Hermit crabs Pagurus 

Horse conchs Busycon 

Horsehair-snakes Gordiacea 

Horseshoe crabs . . . Limulus polyphemus 

Horse mussels Modiolus modiolus 

Hydra-worms young of Nicolea 

Jack-knife clams .... Siliqua , Tagelus 
Jellyfishes . Hydromedusae, Scyphomedusae 


King crabs (British usage) .... Limulus 

Knobbed whelks Busycon carica 

Lady crabs Ovalipes 

Leathery tunicates Stye la 

Leeches Hirudinea 

Little-necked clams Mercenaria 

Limpets Acmaea testudinalis 

Lion's mane (jellyfish) Cyanea 

Lobsters Homarus 

Long-finned squid Loligo pealei 

Long-necked clams Mya arenaria 

Lug worms Arenicola (3 species) 

Mantis shrimp . Hoplocarida or Stomatopoda 

Mermaid's toenail Anomia 

Mites (marine) Halacaridae 

Moon shells Lunatia , Natica, 

Neverita , Polinices 

Moss animals Bryozoa (Entoprocta and 


Mud crabs Xanthidae 

Mud shrimp Callianassa , Upogebia 

Mud snails Nassarius obsoletus 

Mussels Mytilus , Modiolus 

Nassa .... Nassarius (several species) 

Northern corals Astrangia danae 

Northern squid Ilex illacebrosus 

Opossum shrimp Mysidacea 

Oysters Crassostrea 

Oyster crabs Pinnotheres 

Oyster drills Urosalpinx 

Pea crabs Pinnotheres 

Periwinkles .... Littorina (3 species) 

Portuguese Man-of-War Physalia 

Prawns Palaemonetes , etc. 

Purple sea urchins . . Arbacia punctulata 

Quahogs, quohogs Mercenaria 

Rag worms . . . Nereis (several species) 

Razor clams Ens is 

Red-jointed fiddler crabs . Uca minax 
Red sponges . . Microciona prolifera 
Ribbed mussels . . . Modiolus demissus 

Ribbon worms Nemertea 

Rock crabs .... Cancer (2 species) 

Rock slaters Liqia 

Sand crabs Emerita talpoida 

Sand dollars . . Echinarachnius parma 

Sand moles Emerita talpoida 

Sand shrimp Crangon 

Scale worms . Polynoidae, Sigalionidae 

Scallops Aequipecten , Pecten 

Scud . . various gammarid Amphipoda 

Sea anemones Actinaria 

Sea clams .... Spisula solidissima 
Sea cucumbers Holothuroidea 

Sea egg . . . sea urchins especially 

. . Strongvlocentrotus drobachiensis 

Sea float Velella mutica 

Sea gooseberry Pleurobrachia 

Sea lettuce ulva 

Sea mice A phroditidae 

Sea peach Boltenia , Stye la 

Sea pens Pennatulacea 

Sea pork Amaroucium 

Sea slugs . . . (mostly) Nudibranchia 
Sea squirts . . . Ascidiacea, Tunicata 

Sea stars Asteroidea 

Sea urchins . . (regular) Echinoidea 

Sea walnut Pleurobrachia 

Shipworms Teredo , Bankia 

Short-finned squid . Ilex illacebrosus 

Skeleton shrimp Caprellidae 

Soft corals . . . Alcyonium carneum , 

Soft-shelled clams . . . Mya arenaria 
Soft-shelled crabs 

. . . post-moult Callinectes sapidus 
Spider crabs . . Libinia (2 species) , 


Sponges Porifera 

Square -backed 

fiddler crab . Sesarma reticulatum 

Squid, common Loligo pealei 

Starfishes Asteroidea 

Stony or true coral . Astrangia danae, 

Haliplanella luciae 

Striped anemones 
Sulfur sponges 
Summer squid . 
Surf clams . . 
Tube -worms . . 

Whale lice . 
Whelks . . . 
White shrimp 
Winkles . . 

. I lex illacebrosus 
Spisula solidissima 
. various sedentary 
Cyami dae ( Capr e 1 li dae) 
. . . Thais , Busycon 
Callianassa stimpsoni 
Littorina (3 species) 



The following index is to genera and higher taxa mentioned in this manual. In 
general, older synonyms have not been included except for those of experimental ani- 
mals that have been extensively used at Woods Hole in previous years, and which are 
widespread in the literature. Specific names of animals, names of plants, localities, 
authors, and anatomical terms have not been indexed. Underlined page numbers indicate 
illustrations. For vernacular and non-scientific names, see the glossary starting on 
page 200. 

Abietinaria , 17 
Acanthodesia , 173 
Acanthodoris , 130, 153 

154, 156, 157, 162 
Acaulis , 10, 17 
Acerotisa , 37 
Acmaea , 129, 130, 134, 

139, 142, 143 
Acoela (Opi sthobranchia) 

130, 153 
Acoela (Turbellaria) 31 

Acrothoracica, 91 
Acteon , 130, 135, 139 
Actiniaria, 25 
Aeginella , 110, 111 
Aetea, 172, 176, 182, 183 
Aeolidia, 130, 155, 163 
Aeolis , see Aeolidia and 

other nudibranch syno- 
nyms 162 ff 
Aequipecten , 144, 145 , 147 

Aeverrillia , 172, 175 

176, 180, 181 
Aqlaophamus , 56, _57, 70 
Aiptaisiomorpha , see Ha- 

Alcippe , see Trypetesa . 
Alcyonacea, 25 
Alcyonaria, 25 
Alcyonidium , 171, 176, 180 

Alcyonium , 25 
Alentiana , 75 
Aligena , 150 
Alloeocoela, 31, 32, 33 
Allorchestes , 115 
Almyracuma , 99, 100, 101 

Alpheus , 121 
Amage , 63 
Amaroucium , 194, 196, 197 

Amathia , 168, 169, 172 

176, 180, 181 
Amblyosyllis , 54, _55 , 80 
Ammotrypane , 72 
Ampelisca , 114 
Ampeliscidae, 112, 114 

Ampharete . 63 
Ampharetidae, 52, 53_, 61 

Amphiblestrum . 176, 186 

Amphicteis . 63 
Amphineura, 130, 133 
Amphinome . 63 
Amphinomidae, 63 
Amphipholis . 188, 189, 190 

Amphipoda, 93, 107 ff 
Amphiporus . 41, 43, 44, 45 
Amphi trite . 82, 83 
Amphiura . see Amphipholis 
Ampithoe . 115 
Ampithoidae, 113, 115 
Anachis . 138, 139 
Anadara . 130, 144, 145 

146, 150 
Anaitides see Phyllodoce 

( Anaitides ) 
Anaperus . 31, 32, 34, 35. 
Anasca, 176 
Ancinus , 103, 106, 110 

Anemonia , 26 
Anquinella , 176 
Annelida, 47 ff 
Anomia , 146, 150 
Anomura, 117, 121, 127 
Anopla, 40, 43 
Anoplodactylus , 84, 85 

86, 82 
Anostraca, 88 
Anthomedusae, 19 
Anthozoa, 8, 25 ff 
Anthuridea, 103, 105 
Antinoella , 75 
Aoridae, 115 
Aphanostoma . 31, 32, 34 

Aphrodita , 64 
Aphroditidae, 60, 64 
Aplacophora, 130, 133 
Aplousobranchia, 198 
Aplysia , 130, 135, 139 
Arabella , 58, 59. , 64 
Arabellidae, 58, 59, 62 
Arbacia , 188, 192 

Area, see Anadara 
Archiannelida, 47 
Arcteobia , 75 
Arctica , 150 

Arcuatula , see Modiolus, 151 
Arenicola , 64 
Arenicolidae, 63 
Argulus , 88 
Aricia , see Orbinia 
Aricidea , 56, 5_7, 73, 74 
Armadi llidium , 105 
Asabellides , 63 
Aschelminthes , 46 
Ascidiacea, 193 ff 
Ascophora, 177 
Asellota, 105, 106 
Asellus , 105, 106, 108, 109 
Astacura, 117, 126 
Astarte , 149, 150 
Asterias , 188, 189, 190, 

Asteroidea, 188, 189 
Astranqia , 22, 23., 25 
Asychis , 69 
Aurelia , 21, 22, 22. 
Autolytinae, 79 
Autolytus , 54, 55., 79, 81 
Axiothella , 69 
Balanomorpha, 89 ff 
Balanoqlossus , see Sacco- 

Balanus , 86, 8_7, 89, 90, 91 
Bankia , 144, 145, 147, 150 
Barentsia , 165, 166, 180 

Barnea, see Cyrtopleura 

148, 151 
Batea , 114 
Bateidae, 113, 114 
Bdellonemertea, 40, 43 
Bdelloura , 34, 35., 36 
Beroe , Beroida, 29 
Bicellariella . 173, 176 

182, 183 
Bicidium , 26 
Bittium , 137, 139 
Bivalvia, 130, 146 ff 
Boltenia , 198 

Bopyroidea , see Epicaridea 
Bostrichobranchus , 195 

196, 197, 199 

Systematic Index 


Botryllus , 193, 194, 196 

Bougainvillia , 9, 10, 11 

13, 14, 17, 19 
Bowerbankia , 172, 176, 180 

Brachiodontes , see Modio - 
lus , 151 
Brachyura, 117, 118ff 

Brada , 67 
Branchellion , 47 
Branchiopoda, 88 
Branchiura, 88 
Brania, 54, 5_5, 80, 81 
Bryozoa, see Entoprocta, 

Buccinum , 138, 139, 142 

Bugula , 173, 176, 182, 183 

186, 187 
Bugulopsis, 177, 186, 187 
Busycon , 130, 139, 140 
Byblis , 114 

Caberea, 177, 186, 187 
Caecum , 134, 140 
Calathura , 105 
Callianassa , 118, 124 

125 , 126 
Callinectes , 120, 122 

123 , 126 
Calliopius , 114 
Calliopiidae, 113, 114 
Callipallene , 84, 85, 86 

Callopora , 174, 177, 182 

Calycella , 13, 15, 17 
Calyptoblastea, 8, 10 
Campanularia , 13 , 15, 16 

Campanulariidae, 15 
Campylaspis , 99, 100, 101 
Cancer , 119, 122, 123., 126 
Cancridae, 119 
Capitella , 64 
Capitellidae, 63, 64 
Caprella , 110, 111 
Caprellidea, 112 
Carcinides , see Carcinus , 

Carcinonemertes , 40, 43 
Carcinus , 119, 122, 123 

Cardita . 148, 150 
Caridea, 117, 121 
Carinogammarus , 114 

Car i noma , 42, 43 
Carmia, see Mycale 
Caudina , 192 
Caulorhamphus , 177, 186 

Cavolina , 130 
Cellaria , 177 
Cellepora , 167, 174, 177 

184, 185 
Cellularia , see Bugulop - 
Cephalocarida, 88 
Cephalochordata, 193 
Cephalopoda, 131, 133 
Cerapus , 115 
Cerastoderma , 150 
Cerebratulus , 42, 43,44 

Ceriantharia, 2 5 
Cerianthus , 25 
Cerithiopsis , 137, 140 
Chaetoderma , 130, 133 
Chaetogordius , 47 
Chaetopleura , 130, 133 
Chaetopteridae, 61, 65 
Chaetopterus , 65 
Chaetozona . 54, 55^ 66 
Chalina , see Haliclona 
Cheilostomata, 171 ff 
Chelicerata, 84 ff 
Chelifera, see Tanaida- 

Chelura, 115 
Cheluridae, 115 
Childia, 31, 32, 34, 35 
Chiridotea , 104, 106 

110, 111 
Chlamys , 130, 147, 150 
Chondrophora, 20 
Chone , 56, 57., 76, 77 
Chordata, 193 ff 
Chrysaora , 22, 2Z_, 24 
Chrysoela , see Haliplan- 

Chrysopetalidae, 54, 55 

Chthamalus , 86, 8_7, 89 

90, 91 
Cingula , 136, 140 
Ciona, 195, 196, 197 

Cirolana, 103, 106, 108 

Cirratulidae, 54, 55_, 61 

Cirratulus , 54, 5_5, 66 
Cirripedia, 88, 89 ff 

Cladocera, 88 
Cladohepatica, 153 
Clavidae, 10 
Cleistocarpidae, 21 
Cliona , 2, 3, 5_, 7 
Clione, 130 
Clymenella , 56, 5_7, 69 
Clytia , 13 , 15, 16, 17 

Cnidaria, 8 ff, 21 ff 

24 ff 
Cochlodesma , 148, 151 
Colus, 138, 139, 140 
Conchostraca, 88 
Conopeum , 173, 177, 186 

Copepoda, 88 
Corambella , 154, 156, 157 

Corbula, 149, 150 
Cordylophora , 10, YS_, 17 
Coronadena , 37 
Corophiidae, 113, 115 
Corophium , 115 
Corymorpha , 14, 17 
Corynitis , see Zanclea 
Coryphella , 155, 158 

159 , 163 
Craqo , see Cranqon , 121 
Crangon , 117, 121, 124, 

Cranqonyx , 114 
Craniella , 6, 7 
Craspedacusta , 19 
Crassinella , 148, 150 
Crassostrea , 130, 146 

Cratena, 155, 160, 161 

162, 163 
Craterolophus , 21, 22 

Crepidula , 130, 134,140 
Cribrilina , 173, 177 

184, 185 
Crisia , 170, 175 
Crucibulum , 134, 140 
Cryptosula , 174, 178,184 

Ctenophora, 8, 29 
Ctenostomata, 171 ff 
Cucumaria , 189, 192 
Cumacea, 93, 98 ff, 101 
Cuminqia , 148, 151 
Cumulata, 33 
Cuspidaria , 131 
Cuthona, 155, 158, 159 


2 04 

Systematic Index 

Cyanea , 21 

Cyathura , 103, 105, 108 

Cvclaspis , 99, 100, 101 
Cyclostomata, 170 ff 
Cydippida, 29 
Cylichna , 135, 140 
Cylindroporella , 178, 186 

Cylista , see Haliplanella 
Cymadusa , 115 
Cynthia , see Stye la 
Cyprina , see Arctica 
Cyrtodaria , 151 
Cyrtopleura , 148, 151 
Cystenides , see Pectinaria 

( Cvstenides ) 
Dalyellioida, 32, 33, 34 

Decapoda (Crustacea) , 117f f 
Dendrobeania , 173, 177, 186 

Dendrodoa , 195, 196, 197 

Dendronotacea, 153, 163 
Dendronotus , 153, 155, 163 
Dentalium , 129, 130 
Derocheilocaris , 88 
Dexamine , 115 
Dexaminidae, 113, 115 
Dexiospira , 56, 57 
Diadumine , 26 
Diadumine , see Haliplan - 
Diastylis , 98, 99, 100 

Didemnum , 194, 198 
Dinophilus , 47, 54, 55 
Diopatra , 64, 71, 72 
Diplocirrus , 67 
Dispio , 58, 59, 78, 79 
Disporella , 170, 175, 180 

Dodecaceria , 54, 55_, 66 
Dolichoqlossus , see Sacco - 

Doridacea, 153, 162 
Doris , see Acanthodoris , 

Doryilleidae, 52, _53, 62 

Doto , see Idulia 
Driloneris , 58, 5_9, 64 
Dysponetus , 54, 5_5, 60 
Echinarachnius , 188, 192 
Echinodermata, 188 ff 
Echinoidea, 188, 192 
Ectopleura , 14, 18 

Ectoprocta, 167 ff 
Edotea , 104, 106, 108 

Edwardsia , 22, 2Z_, 26 

27, 29 
Edwardsia , also see 

Elasmopus , 114 
Electra, 173, 177, 182 

Eloactis , 2 7 
Elysia , 130, 154, 162 
Embletonia , 153, 155 

Emerita , 121, 127 
Endeis, 85 
Enipo , 75 
Enopla, 40, 43 
Enoplobranchus , 54, 55 

Ensis , 130, 144, 145 

147, 151 
Entoprocta, 165 
Eolidacea, 153, 163 
Eolidia , Eolis , see Aeo - 

lidia and other syn- 
onyms in nudibranch 

Epicaridea, 103 
Epitomapta , 190, 191 

Epitonium , 136, 140 
Epizoanthus , 25 
Eleutherocarpidae, 21 
Erichsonella , 104, 106 

108, 109 
Erichthonius , 115 
Erythrops , 94, 96, 97 
Esperella , see Mycale 
Eteone, 56, 57., 74, 75 
Eubranchus , 158, 159 

162, 163 
Euchone , 76, 77 
Eucratea , 172, 177, 186 

Eudendrium , 9, 11, 13, 

14, 18 
Eudorella , 99, 100, 101 
Eudorellopsis , 99, 100 

101 , 102 
Eulalia, 56, 57, 74, 75 
Eumida , 56, 5J7, 74, 75 
Eunice , 52, 53_, 66, 67 
Eunicidae, 52, _53, 62 

Eunoe , see Harmothoe 

( Eunoe ) 75 
Euphrosine , 52, 5_3_, 60 


Euphrosinidae, 52, 53 , 

60, 67 
Euplana , 37, 44, 45 
Eupleura, 138, 140, 142 

Eupomatus , see Hydroid - 

es ( Eupomatus ) 
Eurylepta . 37 
Eurypanopeus , 119, 122 

123 , 126 
Eusyllinae, 79, 80 
E usylli s, 54, 55, 80, 81 
Evadne, 88 

Exoqone , 54, 55_, 80, 81 
Exogoninae, 79, 80 
Exosphaeroma , 103, 106 

110, 111 
Fabricia, 56, 5J7, 60, 76 

Facelina , 155, 160, 161 

Filograna , 56, 5_7, 77 
Fiona , 155, 160, _16_1, 163 
Flabellifera, 103, 106 
Flabelliqera , 67 
Flabelligeridae, 52, 53. 

60, 67 
Flustra , see Flustrell - 

Flustrella , see Flustrel - 

Flustrellidra , 171, 176 

180, 181 
Galyinia , see Eubranchus 
Gammaridae, 113, 114 
Gammaridea, 112 ff 
Gammarus , 114 
Gastropoda, 130, 134ff 

153 ff 
Gattyana , 75 
Gemellaria , see Eucrat - 
Gemma , 149, 151 
Gemmaria , see Zanclea 
Glycera , 58, 5_9, 67, 68 
Glyceridae, 58, 59^, 62 

Gnathiidea, 103 
Gnesioceros , 37, 44, 45 
Golfinqia , 44, 45., 46 
Goniada , 58, 59., 68 
Goniadella , 68 
Goniadidae, 58, 59, 62 

Gon ion emus , 19 
Gonothyraea , 15, 16, 18 
Gordiacea , 45, 46 
Gorgonocephalus , 189 

Systematic Index 


Gouldia , see Crassinella 

" Grantia " , see Scypha 

Grapsidae, 120 

Gunda , see Procerodes 

Gymnoblastea, 9, 10 

Gymnolaemata, 175 

Gyptis , 68 

Gyratrix , 32, 33, 34, 3_5 

Halacaridae, 84 

Halcampa , 27 

Halecium , 10, 13, 15, 18 

Halichondria , 3, 5_, 7 

Haliclona , 3, 5., 6 

Haliclystus , 21, 22, 23 

Haliplanella , 26, 27 

Haloclava , 22, 23, 26, 27 

Halocynthia , 198 

Haminoea, 130, 135, 140 

Haploscoloplos , see Scolo - 

Haplota , 174, 178, 186, 187 
Harmothoe , 52, 5_3, 75, 76 
Harmothoe ( Eunoe ) , 75 
Harmothoe ( Hermadion ) , 75 
Harmothoe ( Lagisca ) , 76 
Haustoriidae, 113, 114 
Hedylopsis , 130 
Hemichordata, 193 
Henricia , 188, 189 
Hermadion , see Harmothoe 

( Hermadion ) , 75 
Hesionidae, 56, 57, 62, 68 
Heterocrypta , 118, 122, 123 

Heteromastus , 64 
Heteromysis , 94, 95, 96, 97 
Heteronemertea, 40, 41, 43 
"heteronereids " , 70, 71 
"heterosyllids", 79 
Hiatella , 151 
Hippa , see Emerita 
Hippodiplosia , 174, 178 
Hippolyte , 117, 121, 124 

Hipponoe , 63 
Hippoporina , 174, 178, 182 

Hippothoa , 167, 174, 178 

184, 185 
Hirudinea, 47 
Holohepatica, 153 
Holothuroidea, 189, 192 
Homarus , 117, 126 
Hoplocarida, 127 
Hoplonemertea, 40, 43 
Hoploplana , 3 7 
Hutchinsoniella , 88 
Hyale , 115 
Hyalella , 115 

Hyalinoecia , 71, 72 
Hydractinia , 9, 10, 11 

13, 18 
Hydractiniidae , 11 
Hydrobia , 136, 140 
Hydroida, 8, 19 
Hydroides ( Eupomatus ) 

56, 57_, 77 
Hydromedusae, 19 ff 
Hydrozoa, 8 ff 
Hypaniola , 52, 5_3, 63 
Hyperia , 110, 111 
Hyperiidea, 112 
Ichthyobdella, 47 
Idotea, 104, 106, 108, 109 
Idulia , 153, 155, 163 
Ilex , 133, 144, 145 
I lyanassa , see Nassarius 
Imogine, 38 
Inachidae, 118 
Inqolfiellidea, 112 
lone , 105, 107, 108, 109 
I schnochiton , 133 
Ischyroceridae, 113, 115 
I schyrocerus , 115 
Isodictya , 5_, 6 
Isopoda, 93, 102 ff, 109 

Jaera, 105, 106, 108 

Jassa , 115 

Kalyptorhynchia, 32 , 33 
Lacuna , 130, 137, 140 
Laeospira , 56, 57 
Laetmatonice , 64 
Laevicardium , 150, 151 
Lagisca , see Harmothoe 

( Lagisca ) , 76 
Lamellibranchia, see Bi- 

Lamprops , 98, 102 
Laomedea , see Gonothy - 

Laonice , 79 
Larvacea, 193 
Leanira , 78 
Lecithophora, 33 
Leichone, 69 
Lembos , 115 
Lepadomorpha, 89 ff 
Lepas , 86, 87, 89, 90 

Lepidametria , 75, 76 
Lepidonotus , 52, 5_3, 75 

Lepralia , 167 
Lepralia has several syn- 
onyms; see ectoproct 


Leptocheirus , 115 
Leptochelia , 102, 105 

108, 109 
Leptocuma , 99, 100, 

101 , 102 
Leptomedusae, 19 
" Leptoplana " , 38 
Leptosynapta , 189, 190 

191 , 192 
Leucon , 99, 100, 101 

Leucosolenia , 2 , 5 , 6 
Libinia, 118, 122, 123 

Lichenopora , 170, 175 
Ligia , 105, 106, 110 

Lilieborgiidae , 113 

Limnomedusae, 19 
Limnoria, 103, 106 

108, 109 
Limulus , 84 

Lineus, 42, 43, 44, 45 
Lironeca, 104, 106, 108 

Lissodendoryx , 5_, 6 
Listriella , 114 
Littorina , 130, 137, 140 

142, 143 
Livoneca, see Lironeca 
Lobata, 28 
Loima , 82 , 83 
Loligo , 131, 133, 144 

Loxosoma , 165, 180, 181 
Lucinoma , see Phacoides 

( Lucinoma ) 
Lumbrineridae 58, 5_9^ 62 

Lumbrineris , 58, 5_9, 68 

Lunatia , 136, 140, 142 

Lycastopsis , 70, 71 
Lyonsia , 130, 148, 151 
Lysianassidae, 112, 114 
Lysianopsis , 114 
Lysilla , 83 
Lysiosguilla , see Nanno - 

squilla , 127 
Macoma , 144, 145, 148 

Macros tomum , 32, 33, 34 

Mactra, see Spisula 

and Mulinia 
Madreporaria, 25 
Magelona , 69 


Systematic Index 

Magelonidae, 61, 69 
Maiidae, 118 
Malacobdella , 40, 43, 44 

Maldane , 56, 5_7, 69 
Maldanidae, 56, 5J7_, 62, 69 
Maldanopsis , 56, 5_7, 69 
Margelopsis , 11 
Marphysa , 52, 53_, 66, 67 
Melampus , 130, 134, 141 
Melinna , 63 
Melita , 114 
Meiribranipora , 167, 173 

177, 182, 183 
Membranipora has numerous 

synonyms; consult ecto- 

proct checklist. 
Menipea , see Tricellaria 
Mercenaria , 130, 144, 145 

149, 150, 151 
Mertensia , 29 
Meterythrops , 94, 95, 96 

Metridium, 26, 27 
Michtheimysis , see Mysis 
Microciona , 3, 5_, 6 
Microdeutopus , 115 
Microphthalmus , 68 
Microstomum , 31, 33, 34 

Microhydra , 19 
Microporella , 174, 178 

182, 183 
Micrura , 42, 43 
Mitrella , 137, 141 
Mnemiopsis , 2 9 
Modiolaria , 147, 151 
Modiolus , 144, 145, 147 

Molgula , 195, 196, 197 

Mollusca, 129 ff, 153 ff 
Monocelis , 31, 33, 34, 35 
Monoculoides , 114 
Monoophorum , 32, 33, 34 35 
Montacuta , 150 
Montaqua , see Cratena 
Mucronella , 175, 178, 179 
Mulinia , 149, 151 
Musculus , 147, 151 
Mya , 130, 144, 145, 148 

149, 151 
Mycale , 5_, 6, 7 
Myriochele , 7 3 
Myrmeciplana , 33 
Mysella , 149, 151 
Mysidacea, 93 ff, 97 
Mysidopsis , 94, 95, 96, 97 
Mysis , 94, 95, 96, 97 

Mystacocarida, 88 
Mytilus , 130, 142, 143 

147, 151 
Myxicola , 76, 77 
Nannosquilla , 127 
Narcomedusae, 19 
Nassa, see Nassarius 
Nassarius , 11, 130, 138 

141, 142, 143 
Natica , 136, 141 
Naushonia , 118, 124, 125 

Neanthes , see Nereis 

( Neanthes ) 
Nectonema , 44, 45, 46 
Nematomorpha, 4_5, 46 
Nematostella , 22, £3_ 

26, 27 
Nemertea, 40 ff, 44, 45 
Neomysis , 94, 95, 96, 97 
Neopanope , 119, 122, 12 3 

Neosperiopsis , see Iso - 

Nephropsida, 117, 126 
Nephtyidae, 56, 57., 62 

Nephtys , 56, 5_7, 70 
Neptunea , 139, 141 
Nereidae, 52, 53, 61 

Nereimyra , 68 
Nereis , 48, 49, 52, 53 

70, 71 
Nereis ( Neanthes) , 52 

53, 70, 71 
Nereis ( Nereis ) , 52, 5_3 

70, 71 
Nerinides , see Scolo - 

Nerocila , 104, 106, 108 

Nicolea, 54, 55., 82, 83 
Nicomache , 69 
Ninoe , 58, 59, 68, 69 
Nolella , 171, 176, 180 

Notaspidea, 130, 153 

Nothria , see Onuphis 

( Nothria ) 
Notocirrus , 64 
Notomastus , 64 
Notophyllum , 56, 52 

74, 75 
Notoplana , 37, 38, 44 

Notostraca, 88 
Nucella, see Thais 

Nucula , 142, 143 , 151 

Nuda, 29 

Nudibranchia, 130, 153ff 

Obelia , 9, 13, 15, 16 

18, 19 
Octocorallia , see Alcy - 

Octopus , 129, 131 
Ocypodidae, 120 
Odontosyllis , 54, 55., 80 

Odostomia , 138, 141 
Oedicerotidae, 113, 114 
Oerstedia , 41, 43 
Oligochaeta, 47 
Ommastrephes , see Ilex 

Onchidoris , 153, 154, 162 

Oncousoecia , 175, 186 

Oniscoidea, 105, 106 
Onuphidae, 52, 5_3, 62, 71 
Onuphis , 52, 5_3_, 71, 72 
Onuphis ( Nothria ) , 71 

Onuphis ( Onuphis ) , 72 
Ophelia , 72 
Opheliidae, 63, 72 
Ophioderma , 188, 189 

190, 191 
Ophioglycera , 58, 59 

Ophiopholis , 188, 190 

191 , 192 

Ophiura , 188, 190, 191 

Ophiuroidea, 188, 189 
Opisthandropora, 33 
Op isthobranchia, 130 

153 ff 
Orbinia , 58, 59, 72, 73 
Orbiniidae, 58, 59_, 63 

Orchestia , 115 
Orchomenella , 114 
Ostracoda , 88 
Ostrea , see Crassostrea 
Ototyphlonemertes , 43 
Ovalipes , 120, 122, 123 

Ovatella , 134, 141 
Owenia , 58, 5_9, 73 
Oweniidae, 58, 5_9, 62 

Oxyurostylis , 98, 100 

101 , 102 
Paqurus , 38, 121, 127 

Systematic Index 


Palaemonetes, 117, 121,124 

Phoxocephalus , 114 

125, 126 

Phyllodoce, 56, 57, 74 

Paleonemertea, 40, 41, 43 


Palio, 153, 154, 163 

Phyllodoce (Anaitides) 

Pallene, see Callipallene 

74, 75 

Pandora, 130, 148, 151 

Phyllodocidae, 56, 5_7 

Panopeus, 119, 122, 123 

61, 74 


Physalia, 20 

Paradexiospira , 56, 57 

Physcosoma, 46 

Parahesione, 68 

Pinnixa, 120, 122, 123 

Paramphinome , 63 


Paranaites, 56, 57, 74, 75 

Pinnotheres, 120, 122 

Paraonidae, 56, 57, 63, 73 

123, 127 

Paraonis , 73 , 74 

Pinnotheridae, 120 

Paraphoxus , 114 

Pista, 54, 55, 82, 83 

Parapionosyllis , 81 

Pitar, 144, 145, 150 

Parapolia, 42, 43 


Parasmittina, 175, 178, 184 

Placopecten, 147, 152 


Plaqiostomum, 32 - 35 

Pareurythoe , 63 

Planes, 120, 122, 123 

Parthenopidae, 118 


Peachia, 27 

Planocera, 38 

Pecten, see Aequipecten, 

Platyhelminthes, 30 ff 


Platynereis, 52, 53, 70 

Pectinaria, Pectinariidae 


58, 59, 60 

Pleurobrachia , 29 

Pedicellina, 165, 166, 180 

Pleurobranchaea , 154 


156, 157, 162 

Pelaqia, 22, 21, 24 

Pleurobranchus , 130 

Pelecypoda, see Bivalvia 

Pleustidae, 115 

Pelia, 118, 127 

Plumulariidae, 9, 16 

Peltoqaster, 92 

Podarke, 56, 57, 62, 68 

Pennaria, 8, 10, 13, 18 

Podocoryne , 9, 11, 13 

Pennatula, 25 

18, 19 

Pennatulacea, 24 

Podon , 88 

Peracarida, 93 ff 

Polinices, 136, 141, 142 

Periploma, 148, 151 


Perophora, 193, 194, 196 

"Polybostrichus" , 79 

197, 198 

Polycera, see Palio 

Petaloproctus , 69 

Polychaeta, 48 ff 

Petalosarsia, 98, 100, 101 

Polychoerus, 31, 32, 34 



Petricola, 148, 151 

Polycirrus , 83 

Phacoides , 152 

Polycladida, 30, 36-38 

Phascolion, 44, 45, 46 

44, 45 

Phascolosoma , see Golfin- 

Polydora, 52, 53, 58 

g_ia, 46 

59, 61, 78, 79 

Pherusa, 52, 53, 67 

Polynoidae, 52, 53., 60 

Phialidium, see Clytia 


Philine, 130, 135, 141 

Polyonyx, 121, 122, 123 

Philoscia, 105, 106, 108 



Polyphysia, 77 

Phlebobranchia, 198 

Polyplacophora, 130, 133 

Pholoe, 78 

Polyzoa, see Ectprocta 

Photidae, 113, 115 

Pontogeneiidae, 113, 114 

Photis, 115 

Pontoqeneia, 114 

Phoxichilidium, 85, 86, 87 

Porcellio, 105 

Phoxocephalidae, 113, 114 

Porella, 175, 178 

Porifera, Iff 

Porina, see Cylindropor - 

Porpita , 20 
Potamilla , 56, 57, 76 

Praunus , 94, 95, 96, 97 
Pr axil lei la , 69 
Praxillura , 69 
Prionospio , 78, 79 
Probopyrus , 105, 107 

108, 109 
Procephalothrix , 42, 43 
Procerodes , 34, 3_5, 36 
Proporidae, 31 
Prosobranchia, 130 
Prostheceraeus , 37, 38 

44, 45 
Prostoma , 40, 43 
Prosuberites , 3, 7 
Protobranchia, 130, 146 
Protochordata, 193 ff 
Protula , 77 
Ptilanthura , 105 
Pulmonata, 130 
Pycnogonida, 84, 85 
Pycnoqonum , 85 
Pyqospio , 78, 79 
Pyramidellidacea, 137 
Renilla , 25 
Retusa , 135, 141 
Rhabdocoela, 31, 32, 33 
Rhamphos tome 11a , 175, 178 
Rhithropanopeus , 119, 127 
Rhizocephala, 92 
Rhodine, 69 
Rhynchocoela, 40 ff, 44 

Rochefortia , 149, 151 
Sabella , 56, 57., 76, 77 
Sabellaria , Sabellariidae 

58, 59, 60, 76 
Sabellidae, 50, 56, 57 

60, 76 
Saccoglossus , 193, 196, 197 
" Sacconereis 1 , 1 79 
Sacculina , 92 
Sacoglossa, 130, 153, 162 
Sagartia , 22, 23_, 26, 2 7 
Sagartia , also see Hali - 

Samytha , 63 
Samythella , 63 
Sarsia , 14, 18 
Scalibregma , 52, 53^, 77, 
Scalibregmidae, 52, 53_ 

63, 77 
Scaphopoda, 130 


Systematic Index 

Schizmopora , 174 
Schizomavella , 175, 178 
Schizoporella , 175, 178 

184, 185 
Schizotricha , 9, 10, 13_ 

16, 18 
Scolecolepides , 58, 5_9, 78 

Scolelepis , 58, 59., 78, 79 
Scoloplos , 58, 59, 72, 73 
Scruparia , 172, 177, 178 

182, 183 
Scrupocellaria , 172, 177 

186, 187 
Scyllaea , 153, 154, 156 

157 , 163 
Scypha , 2, 5_, 6 
Scyphacella , 105, 106, 110 

Scyphistoma larva, 21, 23 
Scyphomedusae, 21 ff 
Scyphozoa, 8, 21 ff 
Seila , 137, 141, 142, 143 
Septibranchia, 131 
Seriata, 33 
Serpulidae, 50, 56, 5_7_, 60 

Sertularia , 8, 9, 10, 13. 

17, 18 

Sesarma , 120, 122, 123, 127 
Sigalion , 78 
Sigalionidae, 60, 78 
Siligua , 147, 152 
Siphonophora, 19, 20 
Sipunculoidea, 45, 46 
Smittina , 167, 175, 178 
Smittoidea , 175 
Solaster , 188, 189 
Solemya , 146, 152 
Sphaeroma , 103, 106, 110 

Sphaerosyllis , 54, 55, 80 
Spinther , Spintheridae, 52 

53_, 60 
Spio , 58, 59_, 78, 79 
Spiochaetopterus , 65 
Spionidae, 52, 53_, 58, 59_ 

61, 78 
Spiophanes , 58, 59_, 78, 79 
Spirorbis ( Dexiospira ) , 56 

57_, 77 
Spirorbis ( Laeospira ) , 56 

57_, 77 
Spirorbis ( Paradexiospira ) 

56, 57_, 77 
Spisula , 149, 152 
Squilla , 124, 125 , 127 
Stauromedusae, 8, 21 
Stauronereis , 52, 53 , 66 

Stegophryxus , 105, 107 

108, 109 
Stenolaemata, see Cvclo- 

Stenostomata, see Cyclo- 

Stenothoe , 114 
Stenothoidae, 113, 114 
Stephanosella , see Schi - 
Sternaspidae, Sternaspis 

Sthenelais , 60, 78 
Stolidobranchia, 198 
Stomachetosella , 178,186 

Stomatopoda, 127 
Streblosoma , 83 
Streblospio , 78, 79 
Streptosyllis , 54, 55,80 
Strongylocentrotus , 188 

Stye la, 195, 196,197,198 
Styelopsis , 198 
Stylactis , 11, 18 
Stylochus , 37," 38,44, 45 
Suberites , 5_, 6, 7 
Syllidae, 54, 55_, 62, 79 
Syllides , 80, 81 
Syllinae, 79, 80 
Syllis , 54, 5_5, 80, 81 
Sympleustes , 115 
Syncoelidium , 34, 35_, 36 
Synoicidae, 198 
Sycon , see Scypha 
Syncoryne , see Sarsia 
Tagelus , 147, 152 
Talitridae, 113, 115 
Talorchestia , 115 
Tanaidacea, 93, 102, 105 

Tanais , 102, 105, 108, 

Tanystylum , 84, 85,86,87 
Tegella , 174, 177 
Tellina , 130, 144, 145 

148, 152 
Tentaculata, 29 
Terebella , 82, 83 
Terebellidae, 54, 55., 61 

Terebellides , 54, 5_5, 82 

Tergipes , 158, 159 . 162 

Teredo , 144, 145,147,152 
Tetrastemma , 41, 43 
Thais , 38, 139, 146 
Thalassinidea, 117, 118 


Thaliacea, 193 
Tharyx , 54, .55, 66 
Thelepus , 82, 83 
Thoracica, 89 ff 
Thuiaria , 17, 18 
Thyasira , 144, 145 , 149 

Thyone , 189, 192 
Tmetonyx , 114 
Tornaria larva, 193 
Trachelobdella , 47 
Trachylina, 19 
Trachymedusae, 19 
Travisia , 72 
Tricellaria , 172 
Trichobranchus , 82, 83 
Tricladida, 30, 36 
Triphora , 136, 146 
Triticella , 172, 176 
Trophonia , see Pherusa 
Trypetesa , 89, 91 
Tubulanus , 42, 43 
Tubular ia , 9, 14, 17, 18 

Tubulipora , 175, 186, 187 
Tunicata, 193 ff 
Turbellaria, 30 ff 
Turritopsis , 10, 19 
Typhloplanoida, 32, 33 
Uca , 120, 121, 122, 123 

Umbonula, 179, 186, 187 
Unciola , 115 
Upoqebia , 118, 124, 125 

Urochordata, 193 ff 
Urosalpinx , 38, 130, 139 

Valkeria, 172, 176 
Valvifera, 103, 106 
Velella, 20 

Venericardia , see Cardita 
Venus , see Mercenaria 
Vermicular ia , 134, 146 
Vesicularia , see Triti - 
Victorella , 171, 176, 180 
Volsella , see Modiolus , 151 
Woodsholia , 32, 33, 34, 35_ 
Xanthidae, 119 
Xiphosura , see Limulus , 84 
Xiphosurida, 84 
Yoldia, 146, 152 
Zanclea , 8, 13_, 14, 19 
Zirfaea , 148, 152 
Zoantharia, 25 
Zoanthidea, 25 
Zygeupolia , 42 , 43 
Zygonemertes , 41, 43, 44,